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PROCEEDINGS 

of the 

Biological Society of 
Washington 



VOLUME 109 
1996 



Vol. 109(1) published 16 April 1996 Vol. 109(3) published 9 October 1996 

Vol. 109(2) published 25 June 1996 Vol. 109(4) published 23 December 1996 



WASHINGTON 
PRINTED FOR THE SOCIETY 



EDITOR 
C. Brian Robbins 



ASSOCIATE EDITORS 
Classical Languages Invertebrates 

Frederick M. Bayer Jon L. Norenburg 

Frank D. Ferrari 
Rafael Lemaitre 

Plants Vertebrates 

David B. Lellinger Gary R. Graves 

Insects 
Wayne N. Mathis 



All correspondence should be addressed to the 

Biological Society of Washington, Smithsonian Institution 

Washington, D.C. 20560 



Allen Press Inc. 
Lawrence, Kansas 66044 



OFFICERS AND COUNCIL 

of the 

BIOLOGICAL SOCIETY OF WASHINGTON 

FOR 1996-1997 



OFFICERS 

President 
STEPHEN D. CAIRNS 

President-Elect 
RICHARD P. VARI 

Secretary 
CAROLE C. BALDWIN 

Treasurer 
T. CHAD WALTER 



COUNCIL 

Elected Members 
JOHN FORNSHELL RAFAEL LEMAITEE 

ALFRED L. GARDNER DIANA LIPSCOMB 

SUSAN L. JEWETT JAMES N. NORRIS 



TABLE OF CONTENTS 



Volume 109 



Adrianov, Andrey V. and Robert P. Higgins. Pycnophyes parasanjuanensis, a new ki- 
norhynch (Kinorhyncha: Homalorhagida: Pycnophyidae) from San Juan Island, Wash- 
ington, U.S.A 236-247 

Almany, Glenn R. and Carole C. Baldwin. A new Atlantic species of Acanthemblemaria 

(Teleostei: Blennioidei: Chaenopsidae): Morphology and relationships 419^29 

Alvarez, Fernando, Maria Elena Camacho, and Jose Luis Villalobos. The first species 
of Prionalpheus from the eastern Pacific, and new records of caridean shrimp (Crus- 
tacea: Decapoda: Caridea) from the western coast of Mexico 715-724 

Ashe, James S., Robert M. Timm, and Milton H. Gallardo. Systematics, distribution, 

and host specificity of Edrabius Fauvel (Insecta: Coleoptera: Staphylinidae) 731-743 

Bayer, Frederick M. The Antarctic genus Callozostron and its relationship to Primnoella 

(Octocorallia: Gorgonacea: Primnoidae) 150 

Bayer, Frederick M. Three new species of precious coral (Anthozoa: Gorgonacea, genus 

Corallium) from Pacific waters 205 

Bayer, Frederick M. The gorgonacean genus Arthrogorgia (Octocorallia: Primnoidae) 605-628 

Bravo, Manuel Rafael and Masaaki Murano. Description of Neobathymysis japonica, a 
new genus and species, and revision of the genus Bathymysis with a new species from 
Japan (Crustacea: Mysidacea: Mysidae) 501-511 

Bueno-Soria, Joaquin and Silvia Santiago-Fragoso. Studies in aquatic insects X: De- 
scriptions of five new species of the genus Culoptila Mosely (Trichoptera: Glosso- 
somatidae) from Mexico 446-^5 2 

Chevaldonne, Pierre and Karine Olu. Occurrence of anomuran crabs (Crustacea: Decap- 
oda) in hydrothermal vent and cold-seep communities: a review 286-298 

Child, C. Allan. Additions to the Pycnogonida fauna of Carrie Bow Cay, Belize, middle 

America 5 26-5 3 2 

Child, C. Allan. Pycnogonida of the western Pacific islands, XII. A recent diving survey 

of Okinawa, Ryukyu Islands 533-539 

Child, C. Allan. Pycnogonida of the western Pacific islands, XIII. Collections from 

Indonesia, Melanesia, and Micronesia 540-559 

Child, C. Allan. The Pycnogonida types of William A. Hilton. II. The remaining un- 

described species 677-686 

Child, C. Allan and Michel Segonzac. Sericosura heteroscela and S. cyrtoma, new 
species, and other Pycnogonida from Atlantic and Pacific hydrothermal vents, with 
notes on habitat and environment 664—676 

Dahlgren, Thomas G. Two new species of Dysponetus (Polychaeta: Chrysopetalidae) 

from Italy and Papua New Guinea 575-585 

Damkaer, David M. Copepod taxonomy: Discovery vs. recognition 687-694 

Desbruyeres, Daniel and Lucien Laubier. A new genus and species of ampharetid poly- 
chaete from deep-sea hydrothermal vent community in the Azores triple-junction 
area 248-255 

Ferrari, Frank D. and E. L. Markhaseva. Parkins karenwishnerae, a new genus and 
species of calanoid copepod (Parkiidae, new family) from benthopelagic waters of the 
eastern Pacific Ocean 264-285 

Fukuoka, Kouki and Masaaki Murano. Siriella tuberculum, a new species (Crustacea: 

Mysidacea: Mysidae) from Akajima Island, Ryukyu Islands, Japan 512-516 

Garcia-Garza, Maria Elena, Gabino A. Rodriguez-Almaraz, and Thomas E. Bowman. 
Spelaeomysis villalobosi, a new species of mysidacean from northeastern Mexico 
(Crustacea: Mysidacea) 97-102 

Gelder, Stuart R. Description of a new branchiobdellidan species, with observations on 

three other species, and a key to the genus Pterodrilus (Annelida: Clitellata) 256-263 

Gelder, Stuart R. A review of the taxonomic nomenclature and a checklist of the species 

of the Branchiobdellae (Annelida: Clitellata) 653-663 

Giray, Cem and Gary M. King. Protoglossus graveolens, a new hemichordate (Hemi- 

chordata: Enteropneusta: Harrimanidae) from the northwest Atlantic 430-445 



Graves, Gary R. Hybrid wood warblers, Dendroica striata X Dendroica castanea (Aves: 
Fringillidae: Tribe Parulini) and the diagnostic predictability of avian hybrid 
phenotypes 373-390 

Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 2. Hybrid 

origin of Eriocnemis soderstromi Butler 764—769 

Graves, Gary R. and Nancy L. Newfield. Diagnoses of hybrid hummingbirds (Aves: 
Trochilidae). 1. Characterization of Calypte anna X Stellula calliope and the possible 
effects of egg volume on hybridization potential 755-763 

Handley, Charles O., Jr. New species of mammals from northern South America: Bats 

of the genera Histiotus Gervais and Lasiurus Gray (Chiroptera: Vespertilionidae) 1-9 

Harvey, Alan W. and Elizabeth M. De Santo. On the status of Pachycheles laevidactylus 

Ortmann, 1892 (Crustacea: Decapoda: Porcellanidae) 707-714 

Healy, Brenda. Records of Enchytraeidae (Annelida: Oligochaeta) from west Florida. 
1. Mesenchytraeus, Cogenettia, Bryodrilus, Hemienchytraeus, Henlea, and 
Buchholzia 118-137 

Humes, Arthur G. Orecturus amplus, a new species (Copepoda: Siphonostomatoida: 

Asterocheridae) from an alcyonacean in New Caledonia 112-117 

Karasawa, Hiroaki and Hisayoshi Kato. Dalldorfia Rathbun, 1904 (Crustacea: Decapoda) 

from the Neogene of Japan 44—52 

Kensley, Brian. Systematics and distribution of the genus Calocarides (Crustacea: De- 
capoda : Axiidae) 5 3-69 

Kensley, Brian. A new species of the axiid shrimp genus Acanthaxius from the Carib- 
bean (Crustacea: Decapoda: Thalassinidea) 70-74 

Kensley, Brian and Mary Bursey. Rediscovery of Cymodocella algonense from South 

Africa (Crustacea: Isopoda: Sphaeromatidae) 91-96 

Knapp, Leslie W. Review of the genus Cociella Whitley (Teleostei: Platycephalidae) 

with the description of three new species 17-33 

Komicker, Louis S. and Kenneth G. McKenzie. The adult male of the myodocopid 
ostracode Philomedes cubitum Komicker, 1975,. from the Strait of Magellan (Crusta- 
cea: Ostracoda: Myodocopina) 5 17-525 

Lechapt, Jean-Paul and David W. Kirtley. Bathysabellaria spinifera (Polychaeta: Sa- 
bellariidae), a new species from deep water off New Caledonia, southwest Pacific 
Ocean 560-574 

Lemaitre, Rafael and Darryl L. Felder. A new species of ghost shrimp of the genus 
Sergio Manning & Lemaitre, 1994 (Crustacea: Decapoda: Callianassidae) from the 
Caribbean coast of Colombia 453—463 

Lewis, Julian J. and Thomas E. Bowman. The subterranean asellids of Texas (Crustacea: 

Isopoda: Asellidae) 482-500 

Lips, Karen R. and Jay M. Savage. A new species of rainfrog, Eleutherodactylus phasma 

(Anura: Leptodactylidae), from montane Costa Rica 744—748 

Lucas, Spencer G. and Robert J. Emry. Early record of indricothere (Marrmialia: Per- 

issodactyla: Hyracodontidae) from the Aral Sea region of western Kazakhstan 391-396 

Lucas, Spencer G. and Robert J. Emry. Late Eocene entelodonts (Mammalia: Artio- 

dactyla) from Inner Mongolia, China 397—405 

Lucas, Spencer G., Robert J. Emry, and Robert W. Purdy. Marine fossil shark (Chon- 

drichthyes) from nonmarine Eocene sediments, northeastern Kazakhstan 349-352 

Manning, Raymond B. and Darryl L. Felder. Nannotheres moorei, a new genus and 
species of minute pinnotherid crab from Belize, Caribbean Sea (Crustacea: Decapoda: 
Pinnotheridae) . 311-317 

Mathis, Wayne N. Australian beach flies (Diptera: Canacidae) 326-348 

McLaughlin, Patsy A. and Janet Haig. A new genus for Anapagrides sensu De Saint 
Laurent-Dechance, 1966 (Decapoda: Anomura: Paguridae) and descriptions of four 
new species 75-90 

McLaughlin, Patsy A. and John P. Hoover. A new species of Aniculus Dana (Decapoda: 

Anomura: Diogenidae) from Hawaii 299-305 

Ng, Peter K. L. and Boris Sket. The freshwater crab fauna (Crustacea: Decapoda: 

Brachyura) of the Philippines. IV. On a collection of Parathelphusidae from Bohol _- 695-706 



Ng, Peter K. L. and Peter Trontelj. Daipotamon minos, a new genus and species of 

potamid crab (Crustacea: Decapoda: Brachyura) from a cave in China __. 476-481 

Okuno, Junji. Cinetorhynchus manningi, a new shrimp (Crustacea: Decapoda: Caridea: 

Rhynchocinetidae) from the western Atlantic 725-730 

Pettibone, Marian H. Review of Hermilepidonotus Uschakov, 1974, and two species of 

polynoid polychaetes (Lepidonotinae) 143-149 

Pettibone, Marian H. Revision of the scaleworm genera Acholoe Claparede, Arctonoella 
Buzhinskaja, and Intoshella Darboux (Polychaeta: Polynoidae) with the erection of 
the new subfamily Acholoinae ___ 629-644 

Rodriguez-Alamaraz, Gabino A. and Ernesto Campos. New locality records of fresh- 
water decapods from Mexico (Crustacea: Atyidae, Cambaridae, and Palaemonidae) .. 34-38 

Rossman, Douglas A. Identity and taxonomic status of the Mexican garter snake Tham- 

nophis vicinus Smith, 1942 (Reptilia: Serpentes: Natricidae) 10-16 

Rozbaczylo, Nicolas and Marco A. Mendez. Artacama valparaisiensis, a new species 
of Terebellidae (Annelida: Polychaeta) from subtidal soft bottoms of Valparaiso Bay, 
Chile 138-142 

Rozbaczylo, Nicolas, Elba Canahuire, and F. Patricio Ojeda. Presence of Micronereis 
in Antarctic waters and description of a new species, M. antarctica (Polychaeta: Ne- 
reididae: Notophycinae) _._. _ 645-652 

Sandberg, Lennart. Hermit crabs of the genus Paguristes (Crustacea: Decapoda: Di- 
ogenidae) from the western Atlantic. Part III. Paguristes markhami, a new species 
from the Bahama and Caicos Islands 470^75 

Savage, Jay M., James R. McCranie, and Mario Espinal. A new species of Eleutherodactylus 

from Honduras related to Eleutherodactylus bransfordii (Anura: Leptodactylidae) 366—372 

Scanlin, Megan and Janet W. Reid. A new copepod species from California, U.S.A.: 

Hesperodiaptomus californiensis (Crustacea: Copepoda: Calanoida: Diaptomidae) 103-111 

Sluys, Ronald. Reconsiderations of the species status of some South American Planarians 

(Platyhelminthes : Tricladida: Paludicola) — 229-235 

Smith, Brian E. and Jonathan A. Campbell. The systematic status of Guatemalan pop- 
ulations of snakes allied with Ninia maculata (Reptilia: Colubridae) 749-754 

Springer, Victor G. and Helen K. Larson. Pholidichthys anguis, a new species of pho- 

lodichthyid fish from Northern Territory and western Australia 353-365 

Stark, Bill P. New species of Macrogynoplax (Insecta: Plecoptera: Perlidae) from Peru 

and Guyana 3 1 8-325 

Tavares, Marcos and Rafael Lemaitre. Lonchodactylus messingi, a new genus and spe- 
cies of Cyclodorippidae (Crustacea: Decapoda: Brachyura) from the Bahamas 464-469 

Werding, Bernd. Description of a new porcellanid, Petrolisthes gertrudae from the 

southeastern Caribbean Sea (Crustacea: Decapoda: Procellanidae) - 306-310 

Westheide, Wilfried and Giinter Purschke. Leptonerilla diplocirrata, a new genus and species 

of interstitial polychaetes from the island of Hainan, south China (Nerillidae) 586-590 

Wicksten, Mary K. Neocrangon zacae (Chace, 1937) synonymized with N. resima 
(Rathbun, 1902), and compared with N. communis (Rathbun, 1899) (Decapoda: Car- 
idea: Crangonidae) _ 39^3 

Woodman, Neal. Taxonomic status of the enigmatic Crypt otis avia (Mammalia: Insec- 
tivora: Soricidae), with comments on the distribution of the Colombian small-eared 
shrew, Cryptotis colombiana 409-4 1 8 

Zecchini, Fulvio, Michael Vecchione, and Clyde F. E. Roper. A quantitative comparison 
of hectocotylus morphology between Mediterranean and western Atlantic populations 
of the squid Illex coindetii (Mollusca: Cephalopoda: Oegopsida: Ommastrephidae) .._ 591-599 



INDEX TO NEW TAXA 



Volume 109 

(New taxa are indicated in italics; new combinations designated n.c.) 

CNIDARIA 
Anthozoa 

Arthrogorgia utinomii 619 

Callozostron acanthodes 161 

deplodiadema 161 

Convexella 171 

megelhaenica n.c _ 180 

Corallium kishinouyei 218 

nix 213 

thrinax 206 

KINORHYNCHA 

Pycnophyes parasanjuanensis 237 

ANNELIDA 
Polychaeta 

ACHOLOINAE 629 

Amathys 248 

lutzi 249 

Artacama valpuraisiensis 139 

Bathysabellaria spinifera 561 

Dysponetus bipapillatus _ 576 

macroculatus 581 

Hermilepidonotus helotypus n.c. 146 

Leptonerilla 586 

diplocirrata 587 

Micronereis antarctica 646 

Paractonoella 639 

aphthalma n.c 640 

indica n.c. 639 

Oligochaeta 

CogneXiia floridae 123 

Mesenchytraeus hamiltoni 120 

Branchiobdellida 

Pterodrilus annulatus 257 

PYCNOGONIDA 

Achelia bullosa 541 

Anoplodactylus brochus 549 

Ascorhynchus petilus - 526 

Parapallene virgosa 535 

Sericosura cyrtoma 671 

heteroscela 667 

Tanystylum papuensis 546 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

ARTHROPODA 

Crustacea 

Acanthaxius kirkmilleri 71 

Aniculus hopperae 299 

Bathymysis distincta 504 

Caecidotea bilineata 490 

Calocarides capensis 55 

macphersoni 58 

Cinetorhynchus manningi 725 

Daldorfia nagashimae 46 

Diapotamon 476 

minos 477 

Hesperodiaptomus califomiensis 104 

Laurentia 76 

albatrossae 76 

balli 84 

senticosa 87 

sibogae 81 

Lirceolus hardeni 483 

Lonchodactylus 464 

messingi 466 

Macrogynoplax ^mrj 321 

kanuku 321 

truncata 318 

yupanqui 318 

Nannotheres 311 

moorei 312 

Neobathymysis 501 

japonica 502 

Orecturus amplus 112 

Paguristes markhami 471 

PARKIIDAE 266 

Parkius 266 

karenwishnerae 266 

Petrolisthes gertrudae 306 

Prionalpheus nayaritae 719 

Sergio sulfureus 453 

Siriella tuberculum 512 

Spelaeomysis villalobosi 97 

Sundathelphusa boex 696 

sottoae 701 

urichi 702 

vedeniki 703 

Insecta 

Chaetocanace ^avi/7^5 332 

koongarra 334 

longicauda 334 

Culoptila acaena 448 

azulae 451 

barrerai 448 

denningi 451 

jamapa 446 

Dynomiella australica 336 

Edrabius australis 735 

chilensiformis 737 

grandis 733 



VOLUME 109, NUMBER 4 

Nocticanace australina 

Procanace mcalpinei 

Xanthocanace collessi .. 



329 
331 
345 



HEMICHORDATA 



Protoglossus graveolens 



Acanthemblemaria johnsoni 

Cociella heemstrae 

hutchinsi 



somaliensis __ 
Pholidichthys unguis 



Eleutherodactylus lauraster 
phasma 



CHORDATA 

Pisces 



Amphibia 



438 



420 
27 
31 
29 

354 



366 

744 



Mammalia 



Histiotus humboldti 
Lasiurus atratus 



61H 
I 



HHOCEEDINGS 

OF THE 

BIOLOGICAL SOCIETY 

OF 

WASHINGTON 



VOLUME 109 NUMBER 1 
16 APRIL 1996 



ISSN 0006-324X 



THE BIOLOGICAL SOCIETY OF WASHINGTON 

1996-1997 
Officers 

President: Stephen D. Cairns Secretary: Carole C. Baldwin 

President-elect: Richard P. Vari Treasurer: T. Chad Walter 

Elected Council 

John Fornshell Rafael Lemaitre 

Alfred L. Gardner Diana Lipscomb 

Susan L. Jewett James N. Norris 

Custodian of Publications: Storrs L. Olson 



PROCEEDINGS 

Editor: C. Brian Robbins 

Associate Editors 

Classical Languages: George C. Steyskal Invertebrates: Jon L. Norenburg 

Frank D. Ferrari 
Plants: David B. Lellinger Rafael Lemaitre 

Insects: Wayne N. Mathis Vertebrates: Gary R. Graves 

Membership in the Society is open to anyone who wishes to join. There are no prerequisites. 
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Known office of publication: National Museum of Natural History, Smithsonian Institution, 
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@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 

109(1): 1-9. 1996 

New species of mammals from northern South America: 
Bats of the genera Histiotus Gervais and Lasiu rus Gra y 

(Chiroptera: Vespertilionidael^^-^JVVT HS0/yJ5A> 

Charles O. Handley, Jr. / ^ MAY Hi 1M& 

Division of Mammals, National Museum of NatWal History, 
Smithsonian Institution, Washington, D.C. 20560S^^S^.g^^^|E,8 

Abstract. — Two strikingly differentiated species of northern South American 
Histiotus and Lasiurus have been long known, but until now have remained 
undescribed and unnamed. The new Histiotus (H. humboldti) is a relict species, 
most like H. montanus Philippi & Landbeck, but differing from it and all other 
Histiotus in small size (forearm <47 mm, maxillary toothrow length <5.7 mm), 
fragile skull, and weak dentition. Isolated populations occur at medium ele- 
vations (1500-2200 m) in the Coast Range, in the Merida Andes, and on Cerro 
Neblina in Venezuela, and in the Central and Western Andes and upper Cauca 
Valley in Colombia. The new Lasiurus (L. atratus) is a Guayanan endemic 
known from Venezuela, Suriname, and French Guiana. Medium size (forearm 
44-47 mm, max. toothrow 1. 4.4-5.0 mm) and black wings relate it to L. varius 
Poeppig of Chile and L. castaneus Handley of Panama and Costa Rica; but 
very bright black and white underparts; small antorbital fossa; obsolete lach- 
rymal process, supraorbital ridge, and basial pits; and well-developed mastoid 
process distinguish it from both. 



Mammals and their ectoparasites were 
collected in Venezuela between 1965 and 
1968 by the Smithsonian Venezuelan Pro- 
ject, supported in part by a contract (DA- 
49-MD-2788) of the Medical Research and 
Development Command, Office of the Sur- 
geon General, U.S. Army. Numerous pa- 
pers have described the ectoparasites and 
mammals of the Project. Throughout these 
papers undescribed species of mammals 
have been referred to by alphabetical des- 
ignations. Some of these have been named 
subsequently by Handley & Ferris (1972), 
Handley & Gordon (1980), Handley (1984, 
1987). This paper provides formal descrip- 
tions for insect-eating bats of the genera 
Histiotus Gervais and Lasiurus Gray. 



the directions of Kalko & Handley (1994). 
Cranial measurements were taken with the 
assistance of a binocular microscope, with 
dial calipers reading to 0.1 mm. 

Specimens are deposited in the following 
institutions: American Museum of Natural 
History, New York (AMNH); Estacion 
Bioldgica de Rancho Grande, Maracay 
(EBRG); Field Museum of Natural History, 
Chicago (FMNH); National Museum of 
Natural History, Washington (USNM); 
Universidad Central de Venezuela, Caracas 
(UCV). 

Systematics 

A new species of Leaf-eared bat, 

genus Histiotus Gervais 



Material and Methods 



One of the first bats netted by the Smith- 
sonian Venezuelan Project at its inception 
Measurements. — All measurements used in July 1965, at Los Venados in the Coast 
in this paper are in millimeters and follow Range overlooking Caracas, was a species 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



of Histiotus. This genus had not been found 
previously in Venezuela. In a few weeks we 
caught three more Histiotus at Los Venados 
and at the Hotel Humboldt higher up on the 
mountain. These represented a new species 
that I describe and name here. 

This species has been mentioned in the 
literature as Histiotus montanus colombiae 
Thomas by Tamsitt & Valdivieso (1966) 
from Cauca, Colombia, and as Histiotus 
sp. A by me (Handley 1976) from Distrito 
Federal, Venezuela, and by Gardner (1990) 
from Cerro Neblina, Venezuela. 

Histiotus humboldti, new species 
(Fig. 1, Table 1) 

Synonymy. — 
Histiotus montanus colombiae Tamsitt & 
Valdivieso, 1966:102 (not Histiotus co- 
lombiae Thomas, 1916). 

Holotype.—USNM 370968, adult female 
(lactating); skin, skull, and skeleton; col- 
lected 24 July 1965 by Charles O. Handley, 
Jr. Original number, SVP 00092. 

Type locality. — Los Venados, 4 km 
NNW Caracas, 10°32'N, 66°54'W, 1498 m, 
Distrito Federal, Venezuela. The holotype 
was caught 150 m west of park headquar- 
ters buildings in a mist net set across a Jeep 
trail in second-growth forest with thick un- 
derbrush. This area is classified as LOWER 
MONTANE humid forest (bh-MB) in the 
Holdridge system (Ewel and Madriz 1968). 

Etymology. — This impressive bat is 
named in honor of the great naturalist Al- 
exander von Humboldt who traveled widely 
in Venezuela in 1799 and 1800 and de- 
scribed many Venezuelan mammals (Hersh- 
kovitz 1987). 

Distribution. — Histiotus humboldti has a 
fragmented range, with apparently isolated 
populations in southwestern Colombia (on 
the lower eastern flanks of the Western An- 
des at El Tambo and Quisquio and near the 
head of the Cauca Valley at Popayan), in 
north-central Colombia in the northern part 
of the Central Andes (La Ceja and Pobla- 
do), in the Merida Andes in western Ven- 



ezuela (near San Juan de Lagunillas), in the 
Coast Range in northern Venezuela (Los 
Venados and Pico Avila), and on Cerro 
Neblina in southern Venezuela. Elevational 
range, from 1498 m at Los Venados to 2217 
m at La Ceja. 

Ecology. — Histiotus humboldti is a mon- 
tane species that occurs at medium eleva- 
tions, lower than H. montanus usually is 
found at this latitude. Specimens from the 
Coast Range in northern Venezuela were 
taken in moist, second-growth, evergreen 
forest; fairly tall at Los Venados, low and 
very dense at Hotel Humboldt on Pico Avi- 
la. Three were taken in forest trails and one 
was netted in a livestock pen where trees 
remained, but underbrush had been cleared 
(Handley 1976). On Cerro Neblina H. hum- 
boldti was taken in scrubby tepuyan vege- 
tation in open areas close to rocky sand 
hills (A. L. Gardner, pers. comm.). 

Diagnosis. — Histiotus humboldti can be 
recognized as a Histiotus by its enormous 
ears, plain (unomamented) snout, Eptesi- 
cus-like skull, and vespertilionid dentition. 
It can be distinguished from other Histiotus 
by its small size, delicate rostrum, fragile 
zygomata, inflated braincase, and small, 
weak dentition. 

Description. — Dorsal coloration bright 
tan to brown, darker where blackish hair 
bases show through; underparts buff with 
fuscous hair bases showing through. Ears 
very large (28-32); anterior lobe wide (4.3- 
5.5) and forming a point where it folds; tra- 
gus relatively short and broad (9—1 1 X 3.5- 
4.0); tibia and fingers relatively short (tib. 
17.5-18.8, F2 39.1-40.6, F3 78.3-80.5, F4 
62.6-65.6, F5 56.2-60.0). Skufl (Fig. 1) 
fragile throughout; rostrum short (max. 
toothrow 1. 5.3-5.6), narrow (max. br. 5.9- 
6.1), and shallow; facial profile sharply 
dished; lachrymal ridge strongly developed; 
supraorbital region bulges, but is not ledged; 
braincase and area of postorbital constric- 
tion notably inflated; zygoma fragile, but 
with a large postorbital process; pterygoid 
processes thin and delicate. Teeth small and 



VOLUME 109, NUMBER 1 




Fig. 1. Dorsal, ventral, and lateral views of the skull and lateral view of the mandible of Histiotus humboldu, 
USNM 560627, male, from Cerro Neblina, Amazonas, Venezuela. A. L. Gardner photograph. Scale 7:1. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
Table 1. — Measurements (in millimeters) and mass (in grams) of adult Histiotus humboldti. 





USNM 

370968* 

Los Venados 

Venezuela 

9 


ucv 

J-03208 

Cerro Neblina 

Venezuela 

9 


FMNH 

72340 

La Ceja 

Colombia 

9 


EBRG 

[US370967] 

Los Venados 

Venezuela 

6 


USNM 

370969 

Pico Avila 

Venezuela 

S 


USNM 

370970 

Pico Avila 

Venezuela 

6 


USNM 

560627 

Cerro Neblina 

Venezuela 

i 


FMNH 

86719 

Popoyan 

Colombia 

6 


Total length 


109 


107 





106 


110 


110 


110 





Tail vertebrae 


50 


50 


— 


52 


48 


47 


51 


— 


Hind foot (dry) 


9 


11 


10 


9 


10 


9 


11 


11 


Ear from notch 


30 


28 


— 


32 


30 


31 


29 


— 


Forearm 


45.7 


46.8 


46.8 


45.5 


45.7 


45.5 


46.9 


46.0 


Tibia 


17.5 


18.0 


18.0 


17.5 


18.3 


17.9 


18.8 


18.6 


Calcar 


26.3 


23.0 


— 


26.1 


22.8 


— 


23.8 


— 


Mass 


— 


11.5 


— 


— 


— 


— 


9.5 


— 


Greatest length 


16.4 


16.5 


— 


16.0 


16.3 


16.4 


16.9 


— 


Zygomatic breadth 


9.6 


9.8 


— 


9.2 


9.5 


9.5 


9.4 


— 


Postorbital breadth 


4.7 


4.5 


— 


4.5 


AA 


4.5 


4.5 


— 


Braincase breadth 


8.1 


8.0 


— 


7.8 


8.3 


7.9 


8.0 


— 


Braincase depth 


5.8 


5.8 


— 


5.6 


5.6 


5.8 


6.2 


— 


Maxillary toothrow 


















length 


5.4 


5.5 


5.4 


5.5 


5.3 


5.5 


5.6 


5.4 


Postpalatal length 


6.5 


6.1 


— 


6.3 


6.4 


6.6 


6.6 


— 


Maxillary breadth 


6.0 


6.0 


5.9 


6.0 


5.9 


6.1 


6.0 


6.1 


Canine breadth 


4.2 


4.2 


4.1 


4.2 


4.1 


4.1 


4.3 


4.2 



* Holotype. 



weak, with low cusps; dental formula 2/3- 
1/1-1/2-3/3 X 2 = 32. 

Comparisons. — Histiotus humboldti dif- 
fers in almost every detail from H. macrotis 
Poeppig and H. velatus I. Geoffroy. It most 
resembles H. montanus colombiae, a taxon 
with which it is sympatric in some areas, 
but these species differ in many details. 
Coloration, both dorsally and ventrally, is 
similar in both species. Ears of H. hum- 
boldti are proportionally about the same 
size as those of H. m. colombiae, but the 
anterior lobe is wider and is more pointed 
than in any H. montanus (not as extreme as 
in H. velatus, however). The tragus is short- 
er and broader than in any other Histiotus. 
The tibia is shorter than in any H. montanus 
and finger proportions resemble H. velatus 
more than they do H. m. colombiae. In fact, 
in all finger measurements, H. humboldti is 
smaller or averages smaller than H. m. co- 
lombiae. 

Skulls of Histiotus montanus and H. ma- 
crotus are similar to one another and quite 
different from H. velatus, but the skull of 
H. humboldti is so distinctive, that com- 



pared with it, the skulls of the other three 
species are relatively similar to one another. 
In that comparison the skull of H. hum- 
boldti looks as though it might even rep- 
resent a different genus. In fact it bears a 
strong superficial resemblance to skulls of 
North American Plecotus E. Geoffroy, 
Idionycteris Anthony, and Euderma H. Al- 
len. Compared with the other species of 
Histiotus, the skull of H. humboldti is more 
delicate throughout; braincase and postor- 
bital are much more inflated; rostrum is 
much shallower and narrower; dishing of 
facial profile is much more pronounced; 
lachrymal ridge is more developed, but the 
supraorbital ledge is undeveloped; zygo- 
mata are less flaring and much more fragile; 
pterygoid processes are more fragile; teeth 
are relatively tiny, very weak, and have 
lower cusps. Its relatively delicate skull and 
weak dentition indicate that Histiotus hum- 
boldti must have a softer diet than the other 
species of Histiotus. 

Remarks. — Linares (1973) reported a 
specimen of Histiotus in the Museum Na- 
tional d'Histoire Naturelle, Paris (MNHN), 



VOLUME 109, NUMBER 1 



collected in 1894 by Bricefio Gabaldon, 
near Merida, Venezuela. Linares regarded 
the specimen as quite different from Co- 
lombian and Ecuadorean H. montanus be- 
cause it had smaller canines and upper pre- 
molars, an enlarged postorbital process on 
the zygoma, and a dished facial profile. He 
thought some features of the specimen were 
reminiscent of H. velatus. However, be- 
cause of scant material and inadequate 
knowledge of variation in Histiotus, he ten- 
tatively identified the specimen as H. m. 
colombiae (not typical). 

Some characters that Linares ascribed to 
the Merida specimen are characteristic of 
Histiotus humboldti. However, the suite of 
measurements of the Merida specimen 
place it within the size range of H. m. col- 
ombiae, and show that it is apparently too 
large to be H. humboldti. I conclude that 
Linares was correct in identifying the Ven- 
ezuelan specimen in MNHN as H. montan- 
us. 

Recently Jesus Molinari (pers. comm.) 
has taken Histiotus humboldti (confirmed 
by measurements and photographs of the 
skull) adjacent to an extensive island of xer- 
ic vegetation at Tierra Negra, 1550 m, ca. 
12 km S San Juan de Lagunillas, Estado 
Merida, Venezuela. Thus, in Venezuela, 
Histiotus montanus is known in the Merida 
Andes, while H. humboldti appears to have 
a fragmented distribution at medium ele- 
vations in the Coast Range, in the Merida 
Andes, and on a Guayanan tepui, peripheral 
to the range of H. montanus. A similar re- 
lationship is observed in Colombia. This 
suggests to me that H. humboldti once had 
a more extensive, continuous range in the 
mountains of northern South America that 
has been overrun and fragmented by H. 
montanus. 

Specimens examined. — Colombia: Antio- 
quia: La Ceja, [ca. 2217 m], 2 ale. 
(FMNH); Poblado, [ca. 1600 m], 1 ale. 
(AMNH). Cauca: Popayan, 1750 m, 1 ale. 
(FMNH). Venezuela: Amazonas: Cerro 
Neblina, Camp II, 2.8 km NE Pico Phelps, 
1820 m, 1 skin & skull (USNM), 1 skin & 



skull (UCV). Distrito Federal: Los Vena- 
dos, 4 km NNW Caracas, 1498 m, 2 skin, 
skull & skeleton (USNM); Pico Avila, 5 km 
NNE Caracas, 2092-2101 m, 1 skin, skull 
& skeleton (USMN), 1 skin & skull 
(EBRG). 

References to other specimens. — Colom- 
bia: Cauca: El Tambo, 1800 m, 1 skin & 
skull (Swedish Mus. Nat. Hist., Tamsitt & 
Valdivieso 1966); Quisquio, 1700 m, 1 skin 
& skull (Swedish Mus. Nat. Hist., Tamsitt 
& Valdivieso 1966). Published measure- 
ments (fa. 46.1, 47.2; max. toothrow 1. 5.4, 
5.6) (Tamsitt & Valdivieso 1966) agree with 
Histiotus humboldti. 

A new species of Red Bat, 
genus Lasiurus Gray 

In collections of the Field Museum of 
Natural History, I found a specimen 
(FMNH 93235) of a strikingly beautiful red 
bat representing an undescribed species. It 
was collected in Suriname in 1961 by Harry 
Beatty. A short time later, in the collections 
of the Universidad Central de Venezuela in 
Caracas. I discovered two more specimens 
of the same species that had been collected 
in Bolivar state in Venezuela in 1962 by 
Juhani Ojasti. None of these specimens has 
been mentioned in the literature, but re- 
cently a specimen from French Guiana has 
been reported by Brosset & Charles-Dom- 
inique (1990) and by Masson & Cosson 
(1992). 

Lasiurus atratus, new species 
(Fig. 2, Table 2) 

Synonymy. — 

Lasiurus spec? Brosset & Charles-Domi- 
nique, 1990:543. 

Lasiurus castaneus Masson & Cosson, 
1992:476 (not Lasiurus castaneus Hand- 
ley, 1960). 

Holotype.—pymW 93235, adult male, 
skin and skull, collected 10 Feb 1961 by 
Harry A. Beatty. 

Type locality. — Kaiserberg Airstrip, Zuid 



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Fig. 2. Dorsal, ventral, and lateral views of the skull of the holotype of Lasiurus atratus, FMNH 93235, 
male, from Kaiserberg Airport, Zuid River, Suriname. Scott Steppan photograph. Scale 5.6:1. 



River, Suriname. According to Stephens & 
Traylor (1985), this is a hilly region that 
had undisturbed lowland rainforest at the 
time of Beatty's visit. It is located about 
03°07'N, 56°27'W, at an elevation of about 
278 m. 

Etymology. — The Latin atratus, dressed 



in black, refers to the black wing mem- 
branes, which lack the ornate finger outlines 
that characterize the common red bat {Lasi- 
urus blossevillii) of South and Central 
America. 

Distribution. — Known only from Saiil in 
southern French Guiana, from the type lo- 



VOLUME 109, NUMBER 1 



Table 2. — Measurements (in millimeters) of adult Lasiurus atratus. 









Venezuel 


a 






Suriname 


Fr. Guiana 




El Dorado 




Imataca 




Kaiserberg 


Saiil 
Masson & 




UCV 
5409 

9 


UCV 

5410 

? 


Ochoa 

2587 

9 


Ochoa 

2588 

9 


Ochoa 
3183 

S 


Ochoa 
506 


FMNH 
93235* 


Cosson 
(1992) 


Total length 


112 


116 














_ 


_ 


Tail vertebrae 


53 


57 


— 


— 


— 


— 








Hind foot (dry) 


10 


10 


11 


11 


11 


11 


10 




Ear from notch 


13 


13 


— 


— 


— 


— 


13 





Forearm 


46.9 


46.8 


47.6 


46.0 


45.9 


45.1 


46.1 


45.3 


Tibia 


21.8 


19.5 


20.8 


21.1 


20.1 


20.2 


19.2 


— 


Calcar 


14.1 


13.6 


16.5 


16.5 


13.9 


14.9 


12.7 


— 


Greatest length 


13.2 


13.0 


12.9 


12.7 


12.5 


12.6 


13.0 


12.9 


Zygomatic breadth 


9.9 


10.2 


9.9 


9.6 


9.3 


10.0 


9.4 


9.6 


Postorbital breadth 


4.3 


4.3 


4.1 


4.3 


4.1 


4.1 


4.8 


4.2 


Braincase breadth 


7.9 


7.8 


7.6 


7.9 


7.7 


1.1 


1.1 


7.6 


Braincase depth 


6.4 


6.2 


6.1 


6.1 


6.0 


6.2 


6.5 


6.5 


Maxillary toothrow 


















length 


4.9 


5.0 


4.9 


4.7 


4.4 


4.8 


4.9 


4.8 


Postpalatal length 


5.9 


6.1 


5.7 


5.8 


5.5 


5.7 


5.8 


5.7 


Maxillary breadth 


6.5 


6.8 


6.8 


6.5 


6.1 


6.8 


6.4 


6.6 


Canine breadth 


5.4 


5.7 


5.5 


5.3 


5.0 


5.5 


5.4 


5.4 



cality in southern Suriname, and from east- 
ern Venezuela (Km. 55 on the highway 
southeast of El Dorado, and in the Imataca 
Forest, ca. 28 km E Tumeremo, Bolivar). 
Elevational range, 100 m in Bolivar to 278 
m at Kaiserberg Airstrip. Probably Lasiurus 
atratus is a Guayanan endemic. 

Ecology. — Localities in Suriname and 
Venezuela where this bat was collected in 
1961 and 1962 were in undisturbed lowland 
rainforest. Tropical humid forest (bh-T) in 
the Holdridge classification (Ewel & Mad- 
riz 1968). The specimen from French Gui- 
ana was netted above a small stream on the 
border of heavy tropical humid forest and 
a cultivated clearing (Masson & Cosson 
1992). Specimens taken in the Imataca For- 
est in Venezuela by Jose Ochoa, 1990- 

1992, and by Ochoa and Elisabeth Kalko in 

1993, were netted over water-filled roadside 
ditches in secondary forest. 

Diagnosis. — Lasiurus atratus is charac- 
terized by red dorsal coloration, black face, 
contrasting black and white chest, long 
black wings, medium size, small anteorbital 
pit, obsolete lachrymal process, slightly at- 
tenuated exoccipital process, well-devel- 



oped mastoid process, small median ante- 
rior mesoterygoid process, obsolete basial 
pits, and relatively large molars. 

Description. — A medium-sized Lasiurus 
(greatest length of skull 12.9-13.2) with 
long wings (forearm 45.3-46.9); dorsal col- 
oration bright rufous red, without white or 
black hair tips; median buffy band of hairs 
much wider than black basal and red distal 
bands; face black; chin reddish; throat, 
chest, and belly contrasting black and white 
or brown and white (hairs white tipped, 
with successive black, pale or dark brown, 
and black bands); prominent white humeral 
spot; flanks buffy; wings black or blackish, 
lacking ornamental outlines around fingers 
and forearm; ears tan; interfemoral mem- 
brane furred to or near distal edge. 

Skull (Fig. 2) with conventional shape of 
red bat group; rostrum broad (max. br. 6.1- 
6.8), but very short (max. toothrow 1. 4.4- 
5.0) and shallow, sloping sharply down- 
ward anteriorly; facial profile straight; 
braincase large and globose, tilted up from 
palatal plane; sagittal crest low; lambdoidal 
crest weak and incomplete; mastoid process 
well-developed; exoccipital process trian- 



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gular, with a somewhat attenuated tip; zy- 
goma weak, straight on dorsal edge; an- 
teorbital pit small; lachrymal process and 
supraorbital ridge obsolete; palate short 
(max. toothrow 1. 4.4-5.0) and wide (max. 
br. 6.1-6.8); anterior median process of 
mesopterygoid fossa short and wide; basial 
pits poorly developed. 

Dental formula 1/3-1/1-2/2-3/3 X 2 = 
32; anterior upper premolar tiny, wedged 
between lingual borders of canine and P4; 
molars moderately large; M3 with fully de- 
veloped second commissure. 

Comparisons. — Three species of red bats 
occur in the Guayana Region. They are 
graded in size. Lasiurus egregius Peters is 
large (fa. 50.0, gr. 1. 15.7), L. atratus is me- 
dium (fa. 45.3-46.9, gr. 1. 12.9-13.2), and 
L. blossevillii Lesson & Gemot is small (fa. 
36^1, gr. 1. 11.5-11.9). In the red phase 
(the only phase known in L. egregius and 
L. atratus) dorsal coloration is similar in all 
three — bright rufous red, with a wide buffy 
median band on hairs. The face is black in 
L. atratus; reddish or dusky in the other 
species. Coloration of the underparts varies 
widely in the three species — all red like the 
dorsum in L. egregius; speckled brown or 
gray and buff, moderately differentiated 
from the dorsum in L. blossevillii; sharply 
differentiated black and white in L. atratus. 
Wings are ornate, with fingers and forearm 
outlined with reddish in L. blossevillii; plain 
black in L. atratus and L. egregius. With 
respect to cranial features, L. atratus shares 
characters with L. egregius and L. blosse- 
villii. but it differs from both of these spe- 
cies in having the anteorbital pit small, the 
lachrymal process obsolete, and the supra- 
orbital process poorly developed. 

Like Lasiurus atratus, its geographically 
remote relatives, L. castaneus of Central 
America and L. varius of Chile, have black 
unornamented wings, but otherwise they 
are quite different from it. Both have short- 
er wings and strongly developed lachrymal 
and supraorbital processes and anteorbital 
pit. L. castaneus has totally different col- 
oration — blackish underparts, dark dorsum. 



and narrow median band on dorsal hairs, as 
well as more elevated braincase, well-de- 
veloped basial pits, and much reduced M3. 
Underparts of L. varius are uniform orange- 
buff, and it has much more robust molars; 
cranial characters relate it more closely to 
L. blossevillii. 

Specimens examined. — Suriname: Nick- 
arie: Kaiserberg Airstrip, Zuid River, [ca. 
278 m], 1 skin and skull (FMNH). Vene- 
zuela: Bolivar: Km 55 on highway south of 
El Dorado, 100 m, 2 skins & skulls (UCV); 
Reserva Forestal Imataca (Unit 5), ca. 28 
km E Tumeremo, 140-180 m, 1 skin & 
skull, 1 skin, skull & skeleton, 2 alcoholics 
with skulls, 7 alcoholics (collection of J. 
Ochoa), 2 alcoholics with skulls (USNM). 
Total 16. 

References to other specimens. — French 
Guiana: 4 km N Saul, 03°40'N, 53°13'W 
(Masson & Cosson 1992). As described and 
measured by Masson and Cosson (1992), 
this specimen can be only Lasiurus atratus 
(see Table 2). 

Acknowledgments 

I am grateful to the curators who allowed 
me to borrow specimens and/or study col- 
lections in their care at the American Mu- 
seum of Natural History. British Museum 
of Natural History, Field Museum of Nat- 
ural History, and the Universidad Central 
de Venezuela. I especially thankful to Jesus 
Molinari, Universidad de los Andes, Meri- 
da, who sent me photographs and measure- 
ments of a specimen of Histiotus humboldti 
he collected near Merida; and to Jose 
Ochoa, Asociacion Venezolana para la Con- 
servacion de Areas Naturales (ACOANA), 
Caracas, who made a special effort to cap- 
ture Lasiurus atratus in Venezuela, loaned 
me what he caught, and donated specimens 
to the National Museum of Natural History. 
My thanks go to A. L. Gardner who has 
kindly let me reproduce his photographs of 
the skull of Histiotus humboldti, to Bruce 
Patterson who arranged for Scott Steppan 
to photograph the holotype of Lasiurus 



VOLUME 109, NUMBER 1 



atratus, to George Venable who digitized 
and computer-enhanced the photographs, 
and to Darelyn Handley, who made the ta- 
bles and edited and word-processed the 
manuscript. I appreciate the patience of Al 
Gardner, Karl Koopman, Tom Munroe, and 
Don Wilson who read and commented on 
the manuscript. 

Literature Cited 

Brosset, A., & P. Charles-Dominique. 1990. The bats 
from French Guiana: a taxonomic, faunistic and 
ecological approach. — Mammalia 54(4):509- 
560. 

Ewel, J. J., & A. Madriz. 1968. Zonas de vida de 
Venezuela. Ministerio de Agricultura y Cria. 
Caracas, 265 pp. 

Gardner, A. L. 1990. Two new mammals from south- 
ern Venezuela and comments on the affinities 
of the highland fauna of Cerro de la Neblina. 
Pp. 41 1^24 in K. H. Redford & J. E Eisenberg, 
eds.. Advances in Neotropical mammalogy. 
Sandhill Crane Press, Gainesville, Florida, 614 
pp. 

Handley, C. O., Jr. 1976. Mammals of the Smithson- 
ian Venezuelan Project. — Brigham Young Uni- 
versity Science Bulletin, Biological Series 
22(5): 1-89. 

. 1984. New species of mammals from north- 
ern South America: A long-tongued bat, genus 
Anoura Gray. — Proceedings of the Biological 
Society of Washington 97:513-521. 

. 1987. New species of mammals from north- 
ern South America: Fruit-eating bats, genus Ar- 
tibeus Leach. Pp. 163-172 in B. D. Patterson & 
R. M. Timm, eds.. Studies in Neotropical mam- 
malogy: essays in honor of Philip Hershkov- 



itz. — Fieldiana: Zoology, n.s. no. 39, Field Mu- 
seum of Natural History, Chicago, 506 pp. 

— , & K. C. Ferris. 1972. Descriptions of new 
bats of the genus Vampyrops. — Proceedings of 
the Biological Society of Washington 84:519- 
523. 

— , & L. K. Gordon. 1980. New species of mam- 



mals from northern South America. Mouse pos- 
sums, genus Marmosa Gray. Pp. 65-72 in J. F 
Eisenberg, ed.. Vertebrate ecology in the north- 
ern Neotropics. Smithsonian Institution Press, 
Washington, 271 pp. 

Hershkovitz, P. 1987. A history of the Recent mam- 
malogy of the Neotropical region from 1492 to 
1850. Pp. 11-98 in B. D. Patterson & R. M. 
Timm, eds.. Studies in Neotropical mammalo- 
gy: essays in honor of Philip Hershkovitz. — 
Fieldiana: Zoology, n.s. no. 39, Field Museum 
of Natural History, Chicago, 506 pp. 

Kalko, E. K. V, & C. O. Handley, Jr. 1994. Evolution, 
biogeography, and description of a new species 
of fruit-eating bat, genus Artibeus Leach (1821), 
from Panama. — Zeitschrift fiir Saugetierkunde 
59(5):257-273. 

Linares, O. J. 1973. Presence de I'Oreillard d'Ame- 
rique du sud dans les Andes Venezueliennes 
(Chiropteres, Vespertilionidae). — Mammalia 
37(3):433-438. 

Masson, D., & J. F. Cosson. 1992. Cyttarops alecto 
(Emballonuridae) et Lasiurus castaneus (Ves- 
pertilionidae), Deux chiropteres nouveaux pour 
la Guyane fran9aise. — Mammalia 56(3):475- 
478. 

Stephens, L., & M. A. Traylor, Jr. 1985. Ornitholog- 
ical gazetteer of the Guianas. — Bird Depart- 
ment, Museum of Comparative Zoology, Har- 
vard University, Cambridge, Massachusetts, 
121 pp. 

Tamsitt, J. R., & D. Valdivieso. 1966. Bats from Co- 
lombia in the Swedish Museum of Natural His- 
tory, Stockholm.— Mammalia 30(1):97-104. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1):10-16. 1996 

Identity and taxonomic status of the Mexican garter snake 
Thamnophis vicinus Smith, 1942 (Reptilia: Serpentes: Natricidae) 

Douglas A. Rossman 

Museum of Natural Science, Louisiana State University, 
Baton Rouge, Louisiana 70803, U.S.A. 

Abstract. — The nominal garter snake species Thamnophis vicinus is shown 
to be a localized color pattern morph of the wide-ranging T. cyrtopsis collaris; 
this morph is known to occur only in three populations in the Mexican state 
of Michoacan. 



Thamnophis vicinus was originally de- 
scribed (Smith 1942) from the vicinity of 
Morelia, Michoacan, on the basis of ten 
specimens differing from T. cyrtopsis only 
in lacking a vertebral stripe and in having 
large dorsal spots arranged in three rows 
rather than four. Bogert & Oliver (1945), 
Milstead (1953), Duellman (1961), and 
Webb (1966) concluded that T. vicinus is 
merely a color pattern morph of T. cyrtopsis 
(specifically of T. cyrtopsis collaris in the 
current taxonomy) and not a distinct spe- 
cies. However, Webb (1978) reexamined 
most of the type series of T. vicinus, and 
six additional specimens from other areas 
in Michoacan that combined pattern fea- 
tures of T. vicinus and T. cyrtopsis collaris. 
He concluded that the taxonomic status of 
T. vicinus is uncertain; for that reason, he 
tentatively resurrected T. vicinus as a sep- 
arate species. 

Materials and Methods 

Preserved specimens were borrowed 
from the: American Museum of Natural 
History (AMNH); California Academy of 
Sciences (CAS); private collection of E. A. 
Liner (EAL); Field Museum of Natural His- 
tory (FMNH); University of Kansas Mu- 
seum of Natural History (KU); Los Angeles 
County Museum of Natural History 
(LACM); Louisiana State University Mu- 
seum of Natural Science (LSUMZ); Muse- 



um of Comparative Zoology, Harvard Uni- 
versity (MCZ); Michigan State University 
Museum (MSU); Texas Cooperative Wild- 
life Collection, Texas A&M University 
(TCWC); Florida Museum of Natural His- 
tory, University of Florida (UF); University 
of Illinois Museum of Natural History 
(UIMNH); University of Michigan Muse- 
um of Zoology (UMMZ); and Collection of 
Vertebrates, University of Texas at Arling- 
ton (UTA). Four standard characters (num- 
bers of ventrals, subcaudals, and maxillary 
teeth; relative tail length) were recorded; 
the results are summarized in Table 1 . Sev- 
eral aspects of color pattern also were noted 
and are detailed under Results and Discus- 
sion. 

Specimens examined include: Colima, 
LSUMZ 7846; Durango, MSU 4434-36; 
Guanajuato, CAS 5848; Guerrero, LACM 
130112; Hidalgo, MCZ 11432, UMMZ 
99085; JaHsco, MSU 9801, 9803 (2 spec); 
Michoacan, FMNH 37116-22, 39058-61, 
100098 (holotype of T vicinus), 126499- 
504, LACM 65252, MCZ 56019, 131014, 
UIMNH 23414, 23435, UMMZ 102510, 
104699, 112537, 112541, 119409-12, 
121546, UTA R-6050-52; Oaxaca, AMNH 
97889, 103091, 103100, 107001, EAL 
1797, LACM 130111-12, LSUMZ 7560, 
UF 11326-27; Queretaro, TCWC 53068; 
Sinaloa, CAS 24077, 24082, KU 40349, 
78923, 83413, LACM 130113, MSU 567; 
Zacatecas, UMMZ 118433. 



VOLUME 109, NUMBER 1 



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Results and Discussion 

I examined 30 specimens from through- 
out Michoacan that could be identified as 
either T. vicinus or T. cyrtopsis collaris. 
Partial or complete supression of the ver- 
tebral light stripe, accompanied by enlarge- 
ment and frequent fusion of two or more 
sets of dark dorsal spots or blotches on at 
least the anterior portion of the body, occurs 
in specimens from three separate areas in 
Michoacan: near Morelia, in the northeast; 
at Tancitaro, in the west-central region; and 
in the Sierra de Coalcoman, in the south- 
west (Fig. 1). The T. vicinus dorsal pattern 
is by no means uniform in these areas. Most 
specimens from the vicinity of Morelia are 
preservative-darkened, hence details of pat- 
tern are sometimes difficult to discern. No 
indication of a vertebral stripe is evident on 
six specimens (FMNH 126499-500, 
126504; MCZ 56019; UIMNH 23435; 
UMMZ 102510), but there appears to be 
one present on UIMNH 23414, as well as 
traces of a fragmented, indistinct one on 
FMNH 100098 (which, ironically, is the 
holotype of T. vicinus). In all examples, the 
anteriormost postnuchal spots are enlarged 
and fused to form transverse blotches that 
extend from the venter to the vertebral row 
and interrupt the lateral stripes. For a vary- 
ing distance thereafter, the lateral blotches 
alternate with the dorsolateral spots, the lat- 
ter being fused transversely across the back 
to form a single, vertebral row of blotches. 
This pattern (Fig. 2) contrasts with that usu- 
ally attributed to T. cyrtopsis collaris, in 
which the vertebral light stripe is distinct, 
both it and the lateral stripes are uninter- 
rupted, there are no transverse bands on the 
neck, and the dorsolateral spots are not 
fused across the back (Fig. 2). 

The vertebral stripe is present on all eight 
specimens from the Sierra de Coalcoman, 
although it is relatively faint in all but one 
(UMMZ 104699). Six specimens from the 
vicinity of Dos Aguas (UMMZ 1 194 11-1 2, 
121546; UTA R-6050-52) have the char- 
acteristic T. vicinus blotch pattern anteriorly 



12 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

102 99 96 




Fig. I. Map of southwestern Mexico showing the range of Thamnophis cyrtopsis (diagonal lines) and the 
areas where animals having a dorsal pattern characteristic of the "vicinus" morph have been collected (black 
blotches). 



(Fig. 3), whereas the two specimens 
(UMMZ 104699, 112537) from farther 
west (toward Coalcoman) have a T. cyrtop- 
sis collaris blotch pattern on the neck (Fig. 
3). 

Finally, a series of 1 1 specimens (FMNH 
37116-22, 39058-61) from Tancitaro, 
which lies about halfway between Morelia 
and the Sierra de Coalcoman, exhibit an ar- 
ray of dorsal patterns. All but one of the 
series (FMNH 37117) have a vertebral stri- 
pe, but it is faint in six of the specimens. 
In terms of the dorsal blotching, four spec- 
imens have a pattern more-or-less typical of 
T. vicinus, four are more characteristic of T. 



cyrtopsis collaris, and three appear to be 
intermediate (Fig. 4). 

None of the authors who have discussed 
T. vicinus has suggested that it differs from 
T. cyrtopsis collaris in any respect other 
than color pattern, although Webb (1978, 
Table 1) showed that specimens referred to 
the former have a lower mean number of 
subcaudals than the latter The specimens I 
examined from within the "range" of T. vi- 
cinus do, indeed, have fewer subcaudals 
(and a proportionally shorter tail) than spec- 
imens of T. cyrtopsis collaris from north- 
western Michoacan (LACM 65252; MCZ 
131014; UMMZ 119409-10) and farther 



VOLUME 109, NUMBER 1 



13 



m^^ 




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-^t./*». X- a~ 



Fig. 2. Upper: Adult male Thamnophis cyrtopsis collaris (UMMZ 102510) from Pino Gordo (= 37 km by 
road W Ciudad Hidalgo), Michoacan, Mexico, showing a dorsal pattern characteristic of the "vicinus" morph. 
Lower: Subadult female Thamnophis cyrtopsis collaris (UMMZ 112541) from Uruapan Parque Nacional, Mi- 
choacan, Mexico, showing a dorsal pattern characteristic of the subspecies. 



14 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 








Fig. 3. Upper: Adult female Thamnophis cyrtopsis collaris (UMMZ 121546) from Dos Aguas, Michoacan, 
Mexico, showing a dorsal blotch pattern characteristic of the "vicinus" morph. Lower: Subadult male Tham- 
nophis cyrtopsis collaris (UMMZ 104699) from the Cerro de los Havillos, near Coalcoman, Michoacan, Mexico, 
showing a dorsal pattern characteristic of the subspecies. 



VOLUME 109, NUMBER 1 



15 





Fig. 4. Upper: Adult female Thamnophis cyrtopsis collaris (FMNH 39059) from Tancitaro, Michoacan, 
Mexico, showing a dorsal blotch pattern characteristic of the "vicinus" morph. Lower: Adult female Thamnophis 
cyrtopsis collaris (FMNH 37122) from Tancitaro, Michoacan, Mexico, showing a dorsal pattern characteristic 
of the subspecies. 



16 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



north in the range of that subspecies, but 
the mean number of subcaudals is not sig- 
nificantly less than that of T. cyrtopsis col- 
laris from Guerrero and Oaxaca to the 
south (Table 1). Interestingly, still farther 
south — in Chiapas and Guatemala — T. cyr- 
topsis collaris has subcaudal counts that ap- 
proach those of the population north of Mi- 
choacan (Webb 1982). 

Because T. vicinus does not differ men- 
surally or meristically from both geograph- 
ically adjacent populations of T. cyrtopsis 
collaris, and differs only inconsistently in a 
few aspects of color pattern, it is apparent 
that T. vicinus represents nothing more than 
a variant pattern morph — and a junior syn- 
onym — of T. cyrtopsis collaris. Analogous 
situations are not uncommon in other spe- 
cies of Thamnophis (see Rossman et al. 
1996). 

Acknowledgments 

I thank the curatorial staffs at the insti- 
tutions mentioned under Materials and 
Methods for the loan of specimens in their 
care. I also am grateful to R. G. Webb for 
providing a set of his raw data on T. cyr- 
topsis collaris, M. Kleiner for photographic 
assistance, J. Boundy for making the map. 



and D. A. Good for constructive criticism 
of the manuscript. 

Literature Cited 

Bogert, C. M., & J. A. Oliver. 1945. A preliminary 
analysis of the herpetofauna of Sonora. — Bul- 
letin of the American Museum of Natural His- 
tory 83(6):297-426. 

Duellman, W. E. 1961. The amphibians and reptiles 
of Michoacan, Mexico. — University of Kansas 
Publications, Museum of Natural History 15(1): 
1-148. 

Milstead, W. W. 1953. Geographic variation in the 
garter snake, Thamnophis cyrtopsis. — Texas 
Journal of Science 5(3):348-379. 

Rossman, D. A., N. B. Ford, & R. A. Seigel. 1996. 
The garter snakes: evolution and ecology. Uni- 
versity of Oklahoma Press, Norman (in press). 

Smith, H. M. 1942. The synonymy of the garter 
snakes {Thamnophis), with notes on Mexican 
and Central American species. — Zoologica 27: 
97-123. 

Webb, R. G. 1966. Resurrected names for Mexican 
populations of black-necked garter snakes, 
Thamnophis cyrtopsis (Kennicott). — Tulane 
Studies in Zoology 13(2):55-70. 

. 1978. A review of the Mexican garter snake 

Thamnophis cyrtopsis postremus Smith with 
comments on Thamnophis vicinus Smith. — 
Contributions in Biology and Geology, Milwau- 
kee Public Museum (19): 1-13. 

. 1982. Taxonomic status of some Neotropical 

garter snakes (genus Thamnophis). — Bulletin of 
the Southern California Academy of Sciences 
81(l):26-40. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1): 17-33. 1996 

Review of the genus Cociella Whitley (Teleostei: Platycephalidae) 
with the description of three new species 

Leslie W. Knapp 

Department of Vertebrate Zoology, National Museum of Natural History, 
Smithsonain Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — The genus Cociella Whitley is diagnosed as having vomerine 
teeth in two discrete patches, teeth in jaws not depressible, diagonal scale rows 
slanting downward above lateral line more numerous than lateral-line scales, 
lateral-line scale pores with a single canal opening to the exterior, iris lappet a 
simple lobe, side of head bicarinate, suborbital ridge bearing one spine under 
middle eye and one spine under rear margin of eye (additional spines may or 
may not be present posterior to eye), and upper preopercular spine distinctly 
longer than lower spines, bearing a small accessory spine on base. Cociella 
punctata (Cuvier) is removed from the synonymy C. crocodilus (Tilesius) and 
the occurence of possible intergrades between C punctata and C crocodilus 
is discussed. Young C punctata appear to utilize the mangroves as nursury 
grounds. Three new species tentatively assigned to Cociella are described, C. 
heemstrai from off the west coast of southern Africa, C somaliensis from off 
Oman and Somalia and C. hutchinsi from the Arafura Sea. A key to the species 
of Cociella is provided that primarily uses differences in arrangement of spines 
on the suborbital ridge, fin ray counts, number of gill rakers, and scale counts 
to separate the species. 



The status of many nominal genera of the 
Platycephalidae remains unclear. Relatively 
little has been published in this regard since 
extensive revision of Japanese flatheads by 
Matsubara & Ochiai (1955). Matsubara & 
Ochai (1955) synonymized Platycephalus 
punctatus (Cuvier in Cuvier & Valenci- 
ennes, 1829) under Platycephalus crocodi- 
lus (Tilesius, 1812), restricting the genus 
Cociella to C crocodilus. The genus Co- 
ciella is here regarded as containing C 
crocodilus, C. punctata, and three new spe- 
cies described below. This paper attempts 
to clarify the limits Cociella and compares 
features between Cociella and Ratabulus. 

The taxonomic significance of the pore 
morphology of the lateral-line scales in the 
Platycephalidae is well documented. Mat- 
subara and Ochiai (1955) found that differ- 
ences in pore structure of the scales were 
useful as generic characters in Cociella, 



Onigocia, Platycephalus and Rogadius. 
This was further corroborated by Hughes 
(1981, 1985) who, in addition to pore struc- 
ture, also included other features of lateral- 
line scale morphology in a comprehensive 
study of the flatheads that utilized scanning 
electron microscopy. The configuration of 
lateral-line scale pores is also considered 
important here. For example, the scale 
pores of Thysanophrys have two canals to 
the exterior and are Y-shaped; those of On- 
igocia also have two canals to the exterior 
and resemble short, stubby Ts. Other flat- 
head nominal genera with two canals in- 
clude Inegocia, Papillolabium, Rogadius, 
Sorsogona and Suggrundus. On the basis of 
having two pore canals, and with the di- 
agonal scale rows above the lateral line be- 
ing equal or nearly so to the number of lat- 
eral-line scales, these genera are removed 
from consideration here. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



A second group of genera has lateral line 
scale pores with one canal to the exterior: 
Cociella; Elates; Grammoplites; Kumoco- 
cius; Leviprora; Platycephalus; and Rata- 
biilus. To separate these from Cociella, it is 
necessary to use additional characters. The 
condition found in Cociella is given in pa- 
rentheses. Vomerine teeth in a single patch 
is diagnostic for Platycephalus (two patch- 
es). A single elongate preopercular spine 
and six dorsal spines is unique to Elates (2- 
3 preopercular spines, 9 dorsal spines). 
Scale rows slanting downward above the 
lateral line closely approximate the number 
of lateral line scales in Kumococius and 
Grammoplites (scale rows are more numer- 
ous than lateral line scales). In Leviprora, 
the type of iris lappet (or umbraculum) on 
the upper surface of the eye is finger-like 
or cirrose (Matsubara & Ochiai 1955:5, Fig. 
2F) and the two upper preopercular spines 
are subequal (iris lappet a simple lobe, up- 
per propercular spine distinctly longest). 
Ratabulus shares many of the diagnostic 
features given here for Cociella but differs 
in having depressible teeth in the jaws, a 
greater number of spines on the suborbital 
ridge beneath the eye, more elongate later- 
al-line scales (Fig. lA) and a smaller and 
more pointed iris lappet. Although there is 
some justification for placing Cociella in 
the synonymy of Ratabulus, such action 
would require additional evidence that is 
not available here. 

Drawings of lateral line scales from the 
species of Cociella, Ratabulus megace- 
phalus and Kumococius rodericiensis ap- 
pear in Fig. lA-G. These ctenoid scales are 
small, rather uniformly rectangular, have 
well-developed radii, and pores that open to 
the exterior through a single slender canal. 
The canal of C. heemstrai is somewhat 
shorter than the canals in other species of 
Cociella. In Fig. lA, Ratabulus megace- 
phalus, the pore and canal is quite similar 
to those of the other species shown but the 
scale is more slender and elongate. The 
scale from Kumococius rodericiensis (Fig. 
IG) appears quite similar to those found in 



Cociella but the spine at the anterior margin 
of the pore is much more robust. 

Methods-Counts follow procedures de- 
scribed by Hubbs & Lagler (1958:19-24) 
with the following exceptions: number of 
diagonal scale rows slanting downward 
(and backward) starting with the row near- 
est the anteriormost lateral line scale and 
ending with row nearest the posterior lateral 
line scale; number of interpelvic scales are 
counted in a straight line between the pelvic 
bases. Measurements (in mm) were taken 
as follows: interorbital width is the least 
bony width between the eyes; orbit diame- 
ter is taken from the lower rear margin of 
the orbit to the base of the preocular spine; 
snout length is the distance from the tip of 
snout to rear base of the preocular spine; 
head length is taken from the tip of the 
snout to the rear margin of the head; and 
standard length is the stright line distance 
from the tip of snout to the rear margin of 
the hypural plate. Counts and measure- 
ments were routinely taken from the left 
side (unless damaged) while gill rakers 
were counted on the right side. The material 
examined was from the following institu- 
tions (abbreviations in parentheses): Acad- 
emy of Natural Sciences of Philadelphia 
(ANSP); American Museum of Natural 
History, New York (AMNH); Australian 
Museum, Sydney (AMS); Bemice P. Bish- 
op Museum, Honolulu (BPBM); California 
Academy of Sciences, San Francisco (CAS, 
SU); Commonwealth Science and Industrial 
Research Organization, Hobart (CSIRO); 
Field Museum of Natural History, Chicago 
(FMNH); Hebrew University, Jerusalem 
(HUJ); Hokkaido University, Hakodate 
(HUMZ); J. L. B. Smith Institute of Ichthy- 
ology, Grahamstown (RUSI); Kanudi Fish- 
eries Research Station, Konedobu, Papua 
New Guinea (KFRS); Museum of Compar- 
ative Zoology, Harvard University (MCZ); 
Museum National d'Histoire Naturelle, Par- 
is (MNHN); Museum fiir Naturkunde der 
Universitat-Humboldt, Beriin (ZMB); Mu- 
seum of Zoology, University of Michigan, 
Ann Arbor (UMMZ); National Museum of 



VOLUME 109, NUMBER 1 



19 




-/^iS»fie /<!4 y 



:^. 



'V^s«^=9e7r-^ 



Fig. 1. Drawings of pored lateral line scales taken from right side (12th scale from front, scale length in 
parentheses) of species of Cociella and related flatheads: A, Ratabulus megacephalus, USNM 329510, 236 mm 
SL, (3 mm); B, Cociella somaliensis, USNM 326300, 216 mm SL, (3 mm); C, Cociella heemstrai, USNM 
326629, 178 mm SL, (2.9 mm); D, Cociella crocodilus, USNM 329509, 261 mm SL, (3.6 mm); E, Cociella 
hutchinsi, USNM 327279, 240 mm SL, (4.3 mm); E Cociella punctata, USNM 327189, 216 mm SL, (4.0 mm); 
G, Kumococius rodericiensis, WAM P26206, 180 mm SL, (4.0 mm). 



Natural History, Washington D.C. (USNM); 
National Natuurhistorische Museum, Lei- 
den (RMNH); Natural History Museum, 
London (BMNH); Natural History Muse- 
um, Los Angeles County (LACM); Natur- 
historisches Museum, Vienna (NMW); 



Northern Territory Museum of Arts & Sci- 
ences, Darwin (NTM); Royal Ontario Mu- 
seum, Toronto (ROM); South African Mu- 
seum, Cape Town (SAM); Western Austra- 
lian Museum, Perth (WAM); Zoologische 
Museum, Universtiteit van Amsterdam 



20 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Interopercular flap (lOP), right side of Co- 
ciella punctata, USNM 329292, 1 1 1 mm SL. 



(ZMA); and Zoologisk Museum, Koben- 
havns Universitet, Copenhagen (ZMUC). 

Code Ha Whitley, 1940 

Cocius Jordan & Hubbs, 1925:286 (type 
species Platycephalus crocodilus Tile- 
sius, 1812, by original designation). 

Cociella Whitley, 1940:243 (substitute for 
Cocius Jordan & Hubbs, preoccupied). 

Diagnosis. — A genus of platycephalid 
fishes characterized by the following: ocular 
papillae absent; iris lappet a simple lobe 
(Figs. 3, 7) or slightly bilobed (latter may be 



an artifact of preservation); anterior nostril 
with an elongate dermal flap posteriorly; in- 
teropercular flap present or absent; pelvic-fin 
rays I, 4+1; anteriormost 1 to 19 lateral-line 
scales bearing small spines; scale rows slant- 
ing downward above lateral line more nu- 
merous than lateral-line scales; lateral-line 
scale pores with a single opening to the ex- 
terior; suborbital ridge bearing one spine un- 
der middle of eye and one spine under rear 
margin of eye, additional spines may or may 
not be present posterior to the eye; a single 
preocular spine; preopercular spines 2 or 3, 
uppermost bearing a small accessory spine 
on base; side of head bicarinate; teeth villi- 
form, caniniform or granular, in broad bands 
on tooth-bearing bones; and vomerine teeth 
in 2 discrete patches. 

Key to the species of Cociella 

lA. Dorsal-fin rays and anal-fin rays usu- 
ally 1 1 ; total number of gill rakers on 
first arch 5-8 2 

IB. Dorsal-fin rays and anal-fin rays usu- 
ally 12; total number of gill rakers on 
first arch 9-18 4 




Fig. 3. Color pattern variation in Cociella crocodilus (After Matsubara & Ochiai, 1955): A, Pattern typical 
of C. crocodilus. Specimen from Kanaiwa, Japan, 285 mm SL; B, Pattern usually found in C. punctata. Specimen 
from the East China Sea, 340 mm SL. 



VOLUME 109, NUMBER 1 



21 



2A. Suborbital ridge with 2 spines below 
eye, several spines behind eye; upper 
preopercular spine long, nearly reach- 
ing to opercular margin; preorbital 

spine slight or lacking 

C. hutchinsi, new species 

2B. Suborbital ridge with 2 spines below 
eye, no spines behind eye; upper preo- 
percular spine shorter, reaching about 
half-way to opercular margin; preor- 
bital spine usually well developed ... 3 

3A. Diagonal scale rows slanting down- 
ward above lateral line 74-91 (X = 
80.6); total gill rakers usually 7 (6-8); 
interopercular flap absent . . C crocodilus. 

3B. Diagonal scale rows slanting down- 
ward above lateral line 60-76 {X — 
67.1); total gill rakers 5-6, usually 6; 
interopercular flap present . . . C. punctata 

4A. Total gill rakers 9-11 {X = 10.0); in- 
terpelvic scale count 16-24 {X = 20.6); 
caudal fin dusky, with small dark spots 
on upper half . .C. heemstrai, new species 

4B. Total gin rakers 12-18 {X = 15.1); in- 
terpelvic scale count 25-35 (X = 29.9); 
caudal fin light, with large dark spots 
and horizontal streaks throughout . . 
C. somaliensis, new species 

Cociella crocodilus 

(Tilesius, 1812) 

Fig. 3A 

Platycephalus crocodilus Tilesius, 1812, pi. 
59, fig. 2 (original description, type lo- 
cality, Nagasaki). — Cuvier in Cuv. & 
Val., 1829:256 (description taken from 
Tilesius). — Beaufort & Briggs, 1962: 
159-161.— Burgess & Axelrod, 1971: 
541, fig. 510. — Burgess & Axelrod, 
1974:1006, fig. 302. 

Platycephalus punctatus. — Giinther, 1880: 
66. 

Platycephalus inermis (not Silurus inermis 
Houttuyn, 1878) Jordan & Evermann, 
1903:361. 

Thysanophrys crocodilus. — Jordan & Rich- 
ardson, 1908:638-640, fig. 4.— Jordan & 
Metz, 1913:54, fig. 49.— Jordan et al., 
1913:286, fig. 235. 

Inegocia crocodilus. — Jordan & Thomp- 
son, 1914:279. 



Cocius crocodilus. — Jordan & Hubbs, 
1925:287. 

Cociella crocodila. — Matsubara & Ochiai, 
1955:87-89, figs. 32, 33 (in part).— 
Anonymous, 1975:214, pi. 208. — Masu- 
da et al., 1975:342, pi. 146, figs. F, G.— 
Kyushin et al. (eds.), 1982:276, fig. 
255.— Masuda et al. (eds.), 1984:322, pi. 
289, figs. C, D. 

Inegocia crocodila. — Tomiyama & Abe, 
1963:69, fig. 201. 

Material examined (63 specimens). — Ja- 
pan: UMMZ 183219 (5, 140-174) Niigita. 
UMMZ 183220 (1, 181) Niigita. SU 23650 
(1, 285) Naoetsu. UMMZ 183218 (2, 240- 
256) Namerikawa. UMMZ 183222 (1, 250) 
Namerikawa. FMNH 104717 (3, 142-204) 
Tokyo. FMNH 71864 (1, 193) Yokohama. 
MCZ 48821 (1, 140) Yenosima. MCZ 
31160 (1, 126) Yenosima. UMMZ 198916 
(1, 178) Toba. FMNH 58790 (1, 142) Toba. 
FMNH 57413 (5, 230-293) Osaka. USNM 
151815 (1, 239) Kobe. NMW 11179 (2, 
332-403) Kobe. ZMB 18778 (2, 251-258) 
Kobe. USNM 10734 (4, 192-216) Wakan- 
oura. USNM 62317-18 (2, 112-175) Hi- 
roshima & Onomichi. UMMZ 183215 (1, 
218) northern Kyushu. CAS 120708 (2, 
241) Nagasaki. SU 13362 (1, 350) Naga- 
saki. ZMUC P80233 (1, 146) Nagasaki. 
Korea, Pusan: UMMZ 183216 (5, 133- 
206). UMMZ 198900 (2, 159-172). USNM 
143416 (2, 126-174). FMNH 55763 (1, 
251). China: SU 31261 (1, 242). ZMA 
112.715 (1, 339) near Chin-huang-tao. 
MCZ 13789 (1, 253) Shanghai. SU 32766 
(1, 218) Ting-hai. USNM 130414 (1, 227) 
Ningpo. USNM 327194 (1, 245) northern 
East China Sea. HUMZ 108647 (1, 308) 
East China Sea. USNM 329509 (1, 261) 
Dongxiang. UMMZ 198909 (1, 88) Fukien. 
Taiwan: CAS 30012 (1, 231) Formosa 
Strait. CAS 15234 (1, 218) Formosa Strait. 
CAS 107973 (1,171) Chilung. Hong Kong: 
ZMUC P80193-194 (2, 80-85). 

Possible intergrades, C. crocodilus X C. 
punctata (5 specimens). — MNHN 05-222 
(1, 170) Bay d' Along, Gulf of Tonkin. SU 



22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Table 1 . — Number of diagonal scale rows slanting downward above lateral line in the species of Cociella. 



53-55 56-58 59-61 62-64 65-67 



-70 71-73 74-76 77-79 80-82 83-85 86-88 89-91 n 



C. crocodihis 














C. heemstrai 


1 


16 


24 


14 


6 


1 


C. hutchinsi 






6 


14 


5 


1 


C. punctata 




2 


1 


16 


29 


34 


C. somaliensis 


2 


12 


11 


7 


2 





li 



17 9 



34 23 



61 


81.0 


62 


60.7 


26 


62.9 


11 


67.9 



34 59.9 



14151 (1, 337) S.W. Kwangtung. HUMZ 
109566 (1, 269) South China Sea, off Sa- 
rawak, Borneo. USNM 32924 (1, 394), 
Penghu Is., Taiwan. FMNH 47490 (1, 54) 
Singapore. 

Description. — (Mean values appear in 
parentheses). Dorsal-fin rays I, VIII, 10-11 
(10.9), usually 11; anal-fin rays 10-12 
(11.0), usually 11; pectoral-fin rays 19-22 
(20.3); pored lateral line scales 51-60 
(54.3), the anteriormost 1-19 (8.2) scales 
bearing weak spines; number of diagonal 
scale rows above lateral line slanting down- 
ward 74-91 (81.0); diagonal scale count 
14-20 (16.7); interpelvic scales 28-46 
(34.4); total gill rakers on first arch 6-8 
(7.1) and branched caudal rays 10-12 
(11.3). Least interorbital width going into 
greatest diameter of orbit 2.3-4.3 (2.9). 
Nape, opercle and cheek behind eye cov- 
ered with ctenoid scales; top of head and 
cheek below eye mostly naked, with a few 
embedded scales. Preorbital spine present; 
a pair of small nasal spines. Preopercular 
spines often two, sometimes three; upper- 
most longest, reaching about half-way to 
opercular margin. Infraorbital ridge usually 
smooth over anterior Va of eye, bearing 5- 
8 small spines posteriorly; suborbital ridge 
with one spine below middle of eye, a sec- 
ond spine below rear margin of eye, no 
spines behind eye. Lateral-line scale shown 
in Fig. ID. Interopercular flap absent. 

Color in alcohol. — Five or six dark bands 
usually crossing back. Dark spots on dorsum 
numerous anteriorly (on head, including up- 
per surface of eye and on anterior trunk 
reaching below lateral line), more scattered 
and primarily above lateral line on posterior 



body. Spinous dorsal fin with broad dusky 
margin, clear area at base; second dorsal fin 
with dark spots on rays; anal-fin membranes 
dusky, rays pale; pectoral fin dusky below, 
with vertical rows of dark spots on upper 
half; pelvic fin dusky; caudal fin dusky, with 
a series of dark blotches or streaks that usu- 
ally form a broad marginal band. 

Remarks. — No type specimens are known 
for Cociella crocodilus. The species is based 
on an inconotype (Tilesius 1812: pi. 59, fig. 
2) and a secondary description based on Ti- 
lesius given by Cuvier {in Cuvier & Valen- 
ciennes, 1829:256). A more complete de- 
scription appears in Matsubara & Ochiai 
(1955:87-89). Several characters effectively 
separating Cociella crocodilus and C punc- 
tata include differences in the number of di- 
agonal scale rows slanting downward above 
the lateral line (Table 1), total numbers of gill 
rakers (Table 2) and presence or absence of 
a preopercular flap (Fig. 2). Most C croco- 
dilus have from 77 to 91 scale rows 0C = 
81) while most C. punctata have 56 to 73 
scale rows {X = 67.9). Nearly all of C. croc- 
odilus have 7 or 8 gill rakers {X = 7.1) while 
most C. punctata have 6 gill rakers or less 
(X = 6.0). The flap is absent in C. crocodilus 
but is usually well-developed in C. punctata 
(Fig. 2). This flap may be partially developed 
in young specimens and may be difficult to 
see in larger specimens that were poorly fixed 
or that have been preserved for many years. 

Matsubara & Ochiai (1955) illustrated 
variation in color pattern between a speci- 
men taken from off Kanaiwa, Ishikawa Pre- 
fecture, Japan (Fig. 3 A) and one from the 
East China Sea (Fig. 3B). In general, large 
dark spots on the dorsum are typical for C. 



VOLUME 109, NUMBER 1 23 

Table 2. — Total number of gill rakers on the first arch in the species of Cociella. 



C. crocodilus 


1 


45 


C. heemstrai 






C. hutchinsi 


15 


3 


C. punctata 


1 84 


5 


C. somaliensis 







14 



36 13 



13 



11 



52 


7.1 


63 


10.0 


18 


6.2 


90 


6.0 


39 


15.1 



crocodilus (Fig. 3A), while C. punctata is 
characterized by having smaller and less 
prominent dark spots (Fig. 3B). 

Intergrades. — Possible intergrades be- 
tween C. crocodilus and C. punctata (Table 
3) include three specimens from the north- 
em South China Sea, one from off Sarawak, 
Borneo and one from Singapore (Fig. 4). 
Interopercular flaps present in some speci- 
mens from this area appear to be less 
strongly developed than is typical for C. 
punctata. A zone of intergradation may ex- 
tend into the East China Sea and other areas 
surrounding the South China Sea and it is 
possible that a few specimens identified by 
me as C. crocodilus or C. punctata are ac- 
tually intergrades. This could account for 
some of the overlap in characters for the two 
species as shown in Tables 1 and 2. Addi- 
tional specimens from the South China Sea 
are needed to clarify relationships between 
the two species. If integration is occurring 
between C. crocodilus and C. punctata, it 
must be very limited as both species appear 
to be maintaining separate character states. 

Distribution. — This species is found 
along the coast of China, Taiwan, Korea 
and southern Japan (Fig. 4). Limited data 
indicate that it is taken by trawling at 
depths from 50 to 90 m and attains a max- 
imum size of about 400 mm SL. 

Cociella punctata 

(Cuvier in Cuvier & Valenciennes, 1829) 

Figs. 2, 3B 

Platycephalus punctatus Cuvier in Cuvier 
& Valenciennes, 1829: 243 (original de- 
scription, type locality, Ceylon, Vanikoro 
Is.).— Gunther, 1 860: 1 80.— Sauvage, 



1875:307, pi. 36, figs. 5, 5a.— Day, 1876: 
277, pi. LX, fig. 3. 

Platycephalus malabaricus Cuvier in Cu- 
vier & Valenciennes, 1829:245 (original 
description, type locality, Mahe). — 
Gunther, 1860 (in part): 181. 

Platycephalus quoyi Bleeker, 1856—1857: 
206 (original description, type locality, 
Ternate and Amboina). 

Platycephalus fasciatus Giinther, 1872:397 
(original description, type locality, Ma- 
nila Bay). 

Thysanophrys quoyi. — Fowler, 1927:289. — 
Herre, 1953:582. 

Platycephalus crocodilus. — Barnard, 1927: 
933 (in part).— Smith, 1950:178 (in 
part). — Beaufort & Briggs, 1962:161- 
162 (in part). — Fourmanoir, 1957:277 (in 
part) — Jones & Kumaran, 1980:644, fig. 
550. 

Suggrundus hunti Fowler, 1937:244, figs. 
249, 250 (original description, type lo- 
cality, Rayong, Siam). 

Grammoplites jacksoni Fowler, 1944:175, 
figs. 25, 26 (original description, type lo- 
cality. New Hebrides). 

Cocius crocodilus. — Herre, 1953:578. 

Thysanophrys punctatus. — Munro, 1955: 
253, fig. 736 (fig. is C. crocodilus, after 
Jordan et al., 1913). 

Cociella quoyi. — Munro, 1967:528-529, 
fig. 993. 

Cociella crocodila. — Matsubara & Ochiai, 
1955:87-89, figs. 32, 33 (in part).— 
Gloerfelt-Tarp & Kailola, 1984:121, figs. 
A, B.— Dor, 1984:89-90.— Knapp, 1984: 
no pagination (fig. is C. crocodilus, after 
Jordan et al., 1913).— Bianchi, 1985a:30 
(fig. is C crocodilus, after Jordan et al.. 



24 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



O 



t- < 






m vo ^ 



O (50 



S c 



-H ^ ^ 






^ ^ S -S =" 



'^ r- 



= x: 



o 


l^ 

rt 




a 


t!r 


a. 


C/l 


U 


o 


^ 


y= 


u 


s 


c 
o 


S 






(4-1 

o 






c 

(U 


O 


, 1 


e 


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r73 


a 


(U 


60 


_o 


Xi 






B 

3 


1 


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z 


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1913).— Bianchi, 1985b:28 (fig. is C. 
crocodilus, after Jordan et al., 1913). — 
Knapp, 1986:483, fig. 155.1.— Baranes & 
Golani, 1993:305, pi. 7, fig. 23. 

Material examined (123 specimens). — 
Syntypes: MNHN 6836 (1, 187) Voyage of 
Peron; MNHN 6851 (1, 209) Trincomalee, 
Ceylon; MNHN 5852 (1, 193) Vanikoro I. 
Taiwan: RUST 38443 (1, 223) Tachi. Oki- 
nawa: USNM 75448 (1, 127) Naha. Phil- 
ippines: BMNH 1872.10.18.117 (1, 190) 
Manila. SU 39846 (1, 55) Manila. SU 9598 
(1, 107) Cavite. SU 39021 (1, 122) Nasug- 
bu. USNM 99762 (1, 210) Leyte I. 
SU39020 (1, 203) Iloilo. SU 27219 (1, 170) 
Culion. SU 27218 (1, 190) Dumaguete. SU 
29758 (1, 103) Dumaguete. CAS 81318 (6, 
37-124) Dumaguete. USNM 329287 (1, 
45) Dumaguete. USNM 329511 (1, 42) 
near Dumaguete. USNM 329512 (1, 79) 
near Dumaguete. SU 29756 (1, 141) Jolo I. 
Palau Islands: CAS 81319 (1, 97) Koror I. 
CAS 83122 (1, 130) Koror I. CAS 83121 
(1, 182) Nardueis I. USNM 329288 (1, 243) 
Nardueis I. CAS 81320 (1, 208) Babelthuap 
I. USNM 329292 (2, 102-111) Babelthuap 
I. South Pacific: AMS 1.17482-006 (1, 78) 
Guadacanal I. (1, 26) New Hebrides Is. Pa- 
pua New Guinea: KFRS F.5629.01 (2, 190- 
210) Port Moresby. CSIRO 1517 (2, 156- 
161) Port Moresby. Indonesia: SU 13756 
(1, 330) Manokwari. USNM 327189 (3, 
62-216) Semei I. USNM 327289 ((4, 70- 
146) Misool I. USNM 325917 (5, 45-99) 
Kepuluan Aru, Borear I. WAM 27697.001 
(1, 145) Tanimbar I. BPBM 19433 (1, 100) 
Ambon I. RMNH 5915 (3, 128-182) Am- 
bon «& Ternate Is. ZMA 112.697 (2, 135- 
136) Obi Is. USNM 264806 (1, 72) Bah. 
USNM 264805 (1, 100) Bali. USNM 
264794 (1, 194) Bah. BMNH 1984.1.1.65 
(1, 270) Bah. NTM S.l 1127-042 (1, 116) 
Bah. NTM S. 10733.014 (1, 195) Bah. Gulf 
of Thailand: CAS 81316 (1, 73) Ko Chang 
I. CAS 81317 (1, 227) Bangkok fish mar- 
ket. USNM 32928 (1, 157) Prachuap Kiri 
Khan. ANSP 62861 (1, 150) Siracha. 
ANSP 68247 (1, 75) Rayong. Singapore: 



VOLUME 109, NUMBER 1 



25 




Fig. 4. Distribution of Cociella crocodilus, C. heemstrai, C. hutchinsi, C. punctata, and C. somaliensis. 



SU 30809 (2, 61-76). NMW 11171 (2, 
146). Malaysia: SU 27794 (1, 174) Sanda- 
kan. ZMUC P80196 (1, 210) Malacca. 
ZMUC P80195 (1, 109) Penang. ZMUC 
P80191 (2, 104-153) Penang. AMS B.5024 
(1, 167) Penang. Ceylon: ROM 1878 (1, 
130). USNM 327285 (1, 355) Colombo. In- 
dia: SU 37202 (1, ?) Andaman Is., Port 
Blair. USNM 327288 (1, 227) Cochin. 
USNM 329285 (1, 209) Cochin. SU 41735 
(2, 136-142) Calicut. AMS B.8128 (1, 205) 
Malabar. MCZ 4287 (1, 151) Carnara. 
NMW 11720 (2, 155-232) Bombay. ANSP 
101419 (1, 180) Bombay. USNM 327284 
(1, 288) near Bombay. Pakistan: LACM 
38126 (1, 222) Karachi. USNM 327286 (1, 
267) near Karachi. AMS B.8130 (1, 241) 
Sind. Iran: USNM 327283 (1, 358) Gulf of 
Oman. Western Indian Ocean: USNM 
326291 (1, 200) Zanzibar. MNHN 6848 (1, 
220) Mahe. AMNH 88086 (1, 170) Mada- 
gascar. USNM 327287 (1, 346) Madagas- 
car. MNHN 1994.505 (1, 275) Madagascar. 
SU 31387 (3, 118-175) Durban. ANSP 
55103 (2, 87-160) Durban. BMNH 
1921.3.1.51 (1, 151) Durban. Gulf of Aden: 
Uncataloged, J. M. Rose 296-004 (1, 180) 
Djibouti. Red Sea: NMW 11718 (1, 170) 



Ghalefca. RMNH 15955 (2, 197-219) Ka- 
maran. NMW 11719 (1, 162) Kamaran. 
BPBM 20382 (1, 195) Suakin. NMW 
11167 (1, 164) Quseir. USNM 326280 (8, 
129-185) Hurghada. HUJ 14019 (2, 303- 
320) Gulf of Aqaba. 

Description. (Mean values appear in pa- 
rentheses). — Dorsal-fin rays IX or I, VIII, 
10-12 (11.0); anal-fin rays 11-12 (11.0); 
pectoral-fin rays 19-22 (20.6), usually 20 
or 21; pored lateral line scales 50-56 (53.7) 
usually 53 or 54, anteriormost 1-16 (5.5) 
bearing small spines; number of diagonal 
scale rows above the lateral line slanting 
downward 56-76 (67.9); diagonal scale 
count 10-17 (13.7); interpelvic scales 21- 
41 (29.1); total gill rakers on first arch 5-7 
(6.0); branched caudal rays 10-26 (11.3). 
Least interorbital width going into greatest 
diameter of orbit 1.8-6.0 (3.5). Nape cov- 
ered with ctenoid scales; top of head, oper- 
cle and cheek bear embedded scales. Pre- 
orbital spine present; a pair of small nasal 
spines. Preopercular spines usually three, 
sometimes two; uppermost longest, reach- 
ing about half-way to opercular margin. In- 
fraorbital ridge usually smooth over ante- 
rior Vi of eye, bearing 5-8 (small spines 



26 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 4. — Young stages of Cociella punctata associated with mangrove habitat. 



Locality 


SL (mm) 


Deptf 


1 of capture (m) 


Bottom type 


Thailand, CAS 81316 


73 




0-0.9 


rocky, sand, mud 


Philippines, USNM 329511 


42 




0-1.0 


sand 


Philippines, USNM 329512 


79 




0-0.5 


sand, silt 


Guadalcanal, AMS 1.17482-006 


78 




7 


7 


Palau, USNM 329292 


102-111 




0-0.9 


mud, sand, gravel, cobbles 


Indonesia, USNM 329517 


45-98 




0-1.0 


9 



posteriorly); suborbital ridge with one spine 
below middle of eye, a second spine below 
rear margin of eye, no spines behind eye. 
Lateral-line scale shown in Fig. IF. Intero- 
percular flap present. 

Color in alcohol. — Five or six dark 
bands usually crossing back. Numerous 
small dark spots on dorsum reaching below 
lateral line, more widely scattered posteri- 
orly. Spinous dorsal fin with broad marginal 
dark band, clear area at base; second dorsal 
fin with dark spots on rays; anal fin inter- 
radial membranes dusky, rays pale; pectoral 
fin dusky on lower half, spotted above (fin 
entirely spotted in some specimens from 
western Indian Ocean); pelvic fin dusky; 
caudal fin variable (entirely dusky in a 
few), usually with a broad dark marginal 
band or series of dark spots and horizontal 
streaks, basal area more or less pale. 

Remarks. — Young specimens of Cociella 
punctata appear to be closely associated 
with mangrove habitat (Table 4). The small- 
est typically have a broad dark saddle 
across the back in the area of the spinous 
dorsal-fin and a narrow saddle near the rear 



of the soft dorsal-fin (Fig. 5). In juvenile to 
adult C punctata, five or six dark bands 
may cross the back (Fig. 3B) or in some, 
the bands may become obscure. A similar 
pattern is also found in juvenile to adult C. 
crocodilus. 

One specimen of C punctata (USNM 
326291) from Zanzibar was found by East 
African Marine Fisheries Organization bi- 
ologists in the stomach contents of a Chwa- 
ka sole (species unknown, recorded as 370 
mm in total length). 

Distribution. — This species is known 
from the Red Sea to South Africa, to Tai- 
wan, Indonesia and to the New Hebrides 
(Fig. 4). As it is common off Port Moresby, 
Papua New Guinea, I would expect C. 
punctata to be found in northern Australian 
waters but, to my knowledge, none have 
been taken to date. Perhaps competition 
from some of Australia's endemic platyce- 
phalids has prevented C punctata from be- 
coming established. It is frequently taken at 
shallow depths by seines or with ichthyo- 
cide and it has been taken by trawl at depths 
from 23-250 m. Two large specimens (303, 




Fig. 5. Young Cociella punctata from mangroves, USNM 329511, 42 mm SL; Negros, Philippines. 



VOLUME 109, NUMBER 1 



27 



320 mm SL) were taken in a trap at 300 m 
in the Gulf of Aqaba (Baranes «& Golani 
1993:305). 



Cociella heemstrai, new species 
Figs. 6, 7 

Platycephalus malabaricus (non Cuvier). — 
Gilchrist & Thompson, 1909:253. — Fow- 
ler, 1925:255. 

Platycephalus tentaculatus (non Rup- 
pell).— Fowler, 1925:255. 

Platycephalus crocodilus (non Tilesius). — 
Barnard, 1927:933 (in part). — Fowler, 
1934:488 (part).— Smith, 1950:378 (in 
part). — Fourmanoir, 1957:274 (in part). 

Cociella sp. — Knapp, 1986:483. 

Material examined. — Holotype, USNM 
326281 (formerly RUSI 13761) (172 mm 
SL) Kenya, 02°38'S, 40°28'E, RA^ Fridtjof 
Nansen, otter trawl, 280 m, 17 Dec 1980, 
Phillip C. Heemstra. 

Paratypes (68): Natal: SAM 10514 (2, 
232-245) Durban Bay. SAM 11877 (1, 
203) Durban Bay. RUSI 1510 (2, 118-157) 
Durban. BMNH 1919.4.1.33 (1, 109) Dur- 
ban. ANSP 54935 (2, 63-136) Durban; 
1931. SU 69736 (1, 118) Durban. SAM 
10035 (1, 216) Natal. RUSI 16492 (2, 135- 
138) Natal; 1914-20. ANSP 77600 (1, 235) 
Tugela R. N of Durban. SAM 11876 (3, 
167-231) South Head, Tugela R., N of Dur- 
ban; 21 Jan 1901. RUSI 36880 (3, 86-198) 
Tugela Bank, N of Durban. RUSI 1510 (2, 
118-157) Durban. Mozambique: RUSI 
10516 (1, 122) Inhaca Island near Louren90 
Marques; Aug 1948. USNM 326296 (1, 
178) Polana near Louren^o Marques; Feb 
1969. RMNH 25144 (1, 172) Louren?© 
Mfirques; 5 Jul 1965. SAM 10822 (1, 215) 
Lourengo Marques; Jun 1920. SAM 26023 
(1, 200) Louren^o Marques; Jun 1920. 
ANSP 77598 (1, 183) Delagoa Bay. USNM 
326297 (1, 175) Delagoa Bay; 6 Feb 1969. 
USNM 326295 (2, 192-210) Delagoa Bay; 
10 Feb 1969. SAM 16725 (1, 202) Delagoa 
Bay. BMNH 1922.2.9.29 (1, 175) Delagoa 
Bay. RUSI 39859 (5, 152-206) Maputo 
Bay (Delagoa Bay); 28 May 1992. USNM 



^ 






:6C6d; 



'•.)-^:<±XPA 



Fig. 6. Holotype of Cociella heemstrai, USNM 
326281, 172 mm SL; Kenya. 



28 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




\ ■[■(.■■ 




Fig. 7. Paratype of Cociella heemstrai, USNM 303749, 180 mm SL; Mozambique Channel. 



326294 (11, 83-144) Delagoa Bay, west 
side; 14 Feb 1969. USNM 326293 (8, 111- 
177) near Beira; 9 Oct 1964. Madagascar: 
MNHN B.2887 (4, 100-140) Nosy Be. 
USNM 326292 (3, 79-166) Northwest 
coast. Bale D'Amboro; 12 Feb 1964. 
USNM 303749 (4, 140-180) 12°42'12"S, 
48°43'06"E; 11 Nov 1988. MNHN 1994.504 
(1, 227) 25°03'S, 47°07'E; 12 Mar 1969. 

Diagnosis. — This species is distin- 
guished from others in the genus Cociella 
in having 9-11 gill rakers on the first arch 
(Table 2) and interpelvic scale count of 1 6- 
24 {X =■ 20.6). It is further distinguished by 
the following combination of characters: 
second dorsal fin and anal fin with 12 rays; 
ratio of snout length divided by interorbital 
width ranging from 2.5-4.4 0L = 3.4); only 
two spines on suborbital ridge below and 
behind eye; and interopercular flap present. 

Description. — (Values for holotype given 
in parentheses). Dorsal-fin rays I, VIII or 
IX (I, VIII), 11-12 (12), usually 12; anal- 
fin rays 11-13 (12), usually 12; pectoral-fin 
rays 19-22 (20), usually 20 or 21; pored 



lateral line scales 52-55 (53), the anterior 
3-19 (10) scales bearing weak spines; scale 
rows above lateral line slanting downward 
and backward 55-69 (64); diagonal scales 
9-15 (12); interpelvic scales 16-24 (20); 
total gill rakers on first arch 9-11 (9), 
branched caudal rays 11-14 (11), usually 
11 or 12. Measurements for the holotype 
and paratypes appear in Table 5. Least in- 
terorbital width going into greatest diameter 
of orbit 1.8-3.4 times (2.2), usually less 
than 2 times in specimens over 215 mm SL. 
Nape, opercle and cheek behind eye cov- 
ered with ctenoid scales; top of head and 
cheek below eye mostly naked, with a few 
embedded scales. Preorbital spine single; a 
pair of small, reclining nasal spines usually 
present; infraorbital ridge smooth anterior- 
ly, bearing 5-7 spines over posterior half of 
eye; suborbital ridge with one spine below 
middle of eye and a second spine near rear 
margin of eye; upper preopercular spine 
reaching nearly to opercular margin. Lat- 
eral-line scale shown in Fig. IC. A narrow, 
elongate interopercular flap present. 



Table 5. — Proportional measurements of Cociella heemstrai expressed in thousandths of standard length. 
Number of specimens given in parentheses after range. 





Holotype 


Paratypes 




Character 


Range 


X 


Standard length (mm) 


172.0 


63.0-245.0 (62) 


149.1 


Head length 


319.8 


301.5-355.0 (62) 


333.1 


Snout length 


95.9 


79.8-111.8 (62) 


98.6 


Orbit diameter (greatest) 


66.9 


60.2-82.9 (62) 


73.4 


Interorbital width (least) 


30.8 


24.0-35.9 (62) 


29.6 


First dorsal spine length 


22.1 


15.0-36.7 (45) 


24.5 


Second dorsal spine length 


130.2 


94.7-166.5 (42) 


124.3 



VOLUME 109, NUMBER 1 



29 




Fig. 8. Holotype of Cociella somaliensis, USNM 326300, 216 mm SL; Somalia. 



Color in alcohol. — Dorsum grayish or 
brownish, sometimes with a few small dark 
spots. Spinous dorsal-fin dusky, with a few 
large dark spots. Soft dorsal-fin bearing nu- 
merous large dark spots. Pelvic fin dusky. 
Pectoral and caudal fins dusky below, with 
small dark spots near upper margin. 

Color in life. — Specimens taken from 
Richards Bay and the Tugela Shelf off Na- 
tal were observed to have a bright yellow 
horizontal bar in the middle of the caudal 
fin. Live specimens seen at the Durban 
Aquarium had dark bands over the back 
and 4 or 5 pairs of white spots along the 
dorsum. The coloration pattern was some- 
what similar to that exhibited by live spec- 
imens of Platycephalus indicus of similar 
size. 

Distribution. — Cociella heemstrai is 
known from Durban, South Africa to Mom- 
basa, Kenya and Madagascar (Fig. 4). Com- 
mon in trawl catches at depths to 280 m., 
it is also taken by seining in shallow estu- 
aries such as Durban Bay. 

Etymology. — Named in honor of Phillip 
C. Heemstra, J. L. B. Smith Institute of Ich- 
thyology, who, over the years, has contrib- 
uted many specimens of flatheads to my 
studies. 

Cociella somaliensis, new species 
Fig. 8 

Material examined. — Holotype, USNM 
326300 (216 mm SL) Western Indian 
Ocean, Somalia, S of Ras Hafun, MA^ 
Beinta Cruise 19, Sta. 13A, 10°13'N, 



5r00'18"E, trawl, 30 m, 8 Feb 1987, Greg 
Small. 

Paratypes (38): Somalia: USNM 326299 
(8, 199-280) same data as holotype. USNM 
302847 (3, 228-290) 10°20'54"N, 51°15'06"E; 
8 Feb 1987. USNM 302846 (1, 271 same 
data as USNM 302847. USNM 326298 (20, 
198-272) 10°07'24"N, 5r31'12"E; 6 Feb 
1987. BMNH 1993.11.5.1-3 (3, 252-236) 
same data as USNM 326298. MNHN 1993- 
0265 (1,213) same data as USNM 326298. 
MNHN 1993-0266 (1, 236) same data as 
USNM 326298. Oman: USNM 326301 (1, 
273) 21°28'N, 59°28'E; 4 Dec 1963. 

Diagnosis. — A species tentatively as- 
signed to Cociella Whitley, it is distin- 
guished from other members of the genus 
in having 12-18 gill rakers on the first arch 
(Table 2) and the ratio of snout length di- 
vided by interorbital width ranging from 
2.0-2.5 {X = 2.2). It is further distinguished 
by the following combination of characters: 
an interpelvic scale count of 25-35 (X = 
29.9); second dorsal fin and anal fin each 
with 12 rays; only two spines on suborbital 
ridge below and behind eye; and interoper- 
cular flap present. 

Description. (Values for holotype given 
in parentheses). — Dorsal-fin rays I, VIII, 
12; anal-fin rays 11-13 (12), usually 12; 
pectoral-fin rays 19-22 (20), usually 20 or 
21; pored lateral line scales 52-55 (53), an- 
terior 6-15 (9) scales bearing weak spines; 
diagonal scale rows above lateral line slant- 
ing downward 54-66 (54); diagonal scale 
count 10-14 (13); interpelvic scales 25-35 



30 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



Table 6. — Proportional measurements of Cociella somaliensis expressed in thousandths of standard length. 
Number of specimens given in parentheses after range. 





Holotype 


Paratypes 




Character 


Range 


X 


Standard length (mm) 


216.0 


199.0-290.0 (38) 


241 A 


Head length 


318.1 


296.9-328.0 (38) 


310.6 


Snout length 


85.6 


84.4-93.8 (38) 


88.2 


Orbit diameter (greatest) 


65.7 


55.4-67.4 (38) 


62.5 


Interorbital width (least) 


38.0 


36.4 44.7 (38) 


39.6 


First dorsal spine length 


24.1 


13.7-28.3 (31) 


20.6 


Second dorsal spine length 


145.8 


119.6-162.0(33) 


146.5 



(30); total gill rakers on first arch 12-18 
(17); branched caudal-fin rays 11-12 (11), 
usually 1 1 . Measurements for holotype and 
paratypes appear in Table 6. Least interor- 
bital width going into greatest diameter of 
orbit 1.3-1.8 (1.7) times. Nape, opercle and 
cheek behind eye covered with ctenoid 
scales; top of head and cheek below eye 
mostly naked, with a few embedded scales. 
Single preocular and preorbital spine; nasal 
spines usually absent; infraorbital ridge 
smooth anteriorly, bearing 5-7 small spines 
over rear half of eye; suborbital ridge with 
one spine below middle of eye and a second 
spine near rear margin of eye; upper preo- 
percular spine curved slightly upward, 
reaching nearly to or slightly beyond oper- 
cular margin. Lateral-line scale shown in 
Fig. IB. A narrow, elongate interopercular 
flap present. 

Color in alcohol. — Dorsum light tan, 
usually with small scattered dark spots. Spi- 
nous dorsal-fin dusky, with large dark spots. 
Soft dorsal-fin pale, with large dark spots. 



Pectoral and pelvic fins dusky. Anal fin 
with pale rays, interradial membrane slight- 
ly dusky. Caudal fin pale, with bold dark 
spots and elongate bars throughout. 

Distribution. — This species is found off 
Somalia and Oman (Fig. 4). It has been 
taken in the trawl catch at depths from 30- 
49 m. 

Etymology. — Named for the country of 
Somalia, off whose shores all but one of the 
known specimens were captured. 

Cociella hutchinsi, new species 
Fig. 9 

IPlatycephalus malabaricus (non Cu- 
vier).— Gunther; 1880:41. 

Suggrundus sp. 1. — Gloerfelt-Tarp & Kail- 
ola; 1984:123, color fig. — Sainsbury, 
Kailola «fe Leyland; 1985:120, color fig. 

Material examined. — Holotype, CSIRO 
1865 (206 mm SL) Australia, Arafura Sea, 
09°16'S, 135°00'E; FRV Soela, trawl, 113 
m, 27 Jun 1981. Paratypes (26): Arafura 




>s;: 



Fig. 9. Holotype of Cociella hutchinsi, CSIRO 1865, 206 mm SL; Arafura Sea (after Sainsbury et al. 1984). 



VOLUME 109, NUMBER 1 



31 



Table 7. — Proportional measurements of Cociella hutchinsi expressed in thousandths of standard length. Num- 
ber of specimens given in parentheses after range. 





Holotype 


Paratypes 




Character 


Range 


X 


Standard length (mm) 


206.0 


142.0-261.0 (23) 


203.4 


Head length 


363.1 


351.4-393.8 (23) 


373.5 


Snout length 


109.7 


107.1-126.3 (23) 


114.1 


Orbit diameter (greatest) 


77.7 


76.2-90.3 (23) 


82.4 


Interorbital width (least) 


20.4 


17.6-25.1 (23) 


20.8 


First dorsal spine length 


14.6 


10.6-20.4 (21) 


15.4 


Second dorsal spine length 


112.6* 


110.0-142.9 (22) 


128.7 



* Indicates broken. 



Sea: BMNH 1879.5.14.222 (1, 192) 9°59'S, 
139°42'E; 10 Sep 1874. AMS 1.27847-020 
(4, 148-259) 10°02'S, 133°58'E; 17 Nov 
1980. AMS 1.21846-002 (2, 254-261) 
09°38'S, 134°02'E; 17 Nov 1980. NTM 
S. 11898-008 (1, 235) 10°18'S, 136°30'E; 
24 Apr 1986. NTM S. 12266-010 (1, 226) 
10°20'S, 134°23'E; 13 Jun 1987. NTM 
S.l 1957-005 (2, 175-230) 10°18'S, 134°08'E; 
16 Oct 1986. NTM S.l 1613-017 (2, 173- 
218) 10°15'S, 136°20'E; 10 Mar 1985. 
USNM 327179 (formerly NTM S.l 1621- 
002) (1, 240) 10°08'S, 136°48'E; 17 Mar 
1985. Timor Sea: CSIRO 2739 (1, 177) 
13°46-48'S, 128°13-14'E; 28 Jun 1980. 
CSIRO 2740 (1, 175) same data as CSIRO 
2739. CSIRO 2741 (1, 165) same data as 
CSIRO 2739. USNM 327190 (3, 159-165) 
same data as CSIRO 2739. WAM P30716- 
001 (4, 142-184) 13°43.5'S, 128°38.6'E; 26 
Dec 1969. USNM 326279 (2, 198-216) 
12°04'S, 127°14-16'E; 29 Jun 1979. 

Diagnosis. — This species is distin- 
guished from other members of the genus 
Cociella in having 3-4 spines on the sub- 
orbital ridge under eye. It is further distin- 
guished by the following combination of 
characters: second dorsal fin and anal fin 
with 11 rays; 6—7 gill rakers on the first 
arch (Table 2); interpelvic scales 23-34 {X 
= 28.1); interopercular flap absent; and ra- 
tio of snout length divided by interorbital 
width ranging from 4.6 to 6.7 {X = 5.6). 

Description. — (Values for holotype given 
in parentheses). Dorsal-fin rays I, VII, I-I, 



VIII (I, VIII), 11; anal-fin rays 11; pectoral- 
fin rays 20-23 (22), usually 21 or 22; pored 
lateral line scales 51-55 (54), the anterior- 
most 2-8 (6) scales bearing weak spines; di- 
agonal scale rows above lateral line slanting 
downward 59-68 (63); diagonal scale count 
11-14 (12); interpelvic scales 23-34 (24); 
total gill rakers on first arch 6—7 (6); 
branched caudal-fin rays 11-12 (12). Mea- 
surements for the holotype and paratypes ap- 
pear in Table 7. Least interorbital width go- 
ing into greatest diameter of orbit 3.2-5.0 
(3.8). Nape, opercle and cheek behind eye 
covered with ctenoid scales, top of head and 
cheek below eye mostly naked, with few 
embedded scales. Preorbital spine slight or 
lacking, a pair of small nasal spines present. 
Infraorbital ridge usually smooth over ante- 
rior Va of eye, bearing 7-10 small spines pos- 
teriorly; suborbital ridge with one spine be- 
low middle of eye, a second spine below 
rear margin of eye and 3-4 spines behind 
eye; upper preopercular spine reaching near- 
ly to opercular margin. Lateral-line scale 
shown in Fig. IE). Interopercular flap ab- 
sent. 

Color in alcohol. — Dorsum brownish; 
venter white with brownish stippling, 
sparse on breast, more evident posteriorly. 
Spinous dorsal fin dusky, with a broad sub- 
marginal black band. Soft dorsal fin pale, 
with small brownish spots on rays. Anal fin 
pale, rays white. Pectoral fin dusky brown, 
with traces of vertical dark bands, lower 
margin white. Pelvic fin with whitish base. 



32 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Stippled with brown, with well-developed 
submarginal dark band. Caudal fin dusky, 
sometimes with a series of streaks forming- 
submarginal dark band. 

Distribution. — This is species is known 
from the Timor and Arafura Seas of Aus- 
tralia. It has been taken by trawling at 
depths from 39-108 m. 

Etymology. — Named in honor of J. Barry 
Hutchins, Western Australian Museum, 
who has provided substantial assistance to 
the author's studies of Australian flatheads. 

Acknowledgments 

I am greatly indebted to the following in- 
dividuals for providing access to specimens 
and/or other assistance: Gerald R. Allen, 
Kunio Amaoka, M. Eric Anderson, Marie 
Louise Bauchot, Adam Ben-Tuvia, Marinus 
Boeseman, Eugenia B. Bohlke, M. Bou- 
gaardt; David C. Catania, Simon Chater, 
Barry Chemoff, Bruce B. Collette, Leonard 
J. V. Compagno, Martine Desoutter, Wil- 
liam N. Eschmeyer, Sean Fennessy, Jerome 
F. Finan, Thomas Gloerfelt-Tarp, Daniel 
Golani, Alastair Graham, Cedric Goliath, 
Karsten E. Hartel, Phillip C. Heemstra, Bar- 
bara Herzig, Jean-Claude Hureau, J. Barry 
Hutchins, Tomio Iwamoto, Robert K. John- 
son, Patricia J. Kailola, Peter Last, Robert 
J. Lavenberg, Anthony D. Lewis, Nigel R. 
Merrett, Douglas W Nelson, Gareth J. Nel- 
son, Jorgen Nielsen, Han Nijssen, John R. 
Paxton, John E. Randall, Billy Ranchod, 
Margaret Rouse, Barry C. Russell, Mark 
Salotti, William G. Saul, Jeffrey A. Seigal, 
William F Smith-Vaniz, Pearl M. Sonoda, 
Arnold Sussumoto, Rex Williams, and 
Richard Winterbottom. The drawings were 
skillfully prepared by Penelope Hollen- 
sworth and Francis W Zweifel. Harold E. 
Dougherty and Rafael Lemaitre took the 
fine photographs. 

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33 



tions of fishes from the coast of Natal (Part 2). 
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the flatheads (Pisces: Platycephalidae). Unpub- 
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obtained in Japan in 1911. — Memoirs Carnegie 
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. 1986. Family No. 155: Platycephalidae. In 

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rica, 482-486, 12 figs., pis. 29-30. 

Kyushin, K., K. Amaoka, K. Nakaya, H. Ida, Y. Tan- 
ino, & T Senta. 1982. Fishes of the South Chi- 
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some new fish names. — Australian Naturalist 
10(8):242-243. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1 ):34-38. 1996 

New locality records of freshwater decapods from Mexico 
(Crustacea: Atyidae, Cambaridae, and Palaemonidae) 

Gabino A. Rodriguez-Almaraz and Ernesto Campos 

(GAR- A) Facultad de Ciencias Biologicas, Universidad Autonoma de Nuevo Leon, A. P. 105-F, 

Ciudad Universitaria, San Nicolas de los Garza, Nuevo Leon, Mexico; 

(EC) Facultad de Ciencias, Universidad Autonoma de Baja California, A. P. 2300, 

Ensenada, Baja California 22800, Mexico 

Abstract. — Collections in the Mexican states of Nayarit, Nuevo Leon, Ta- 
maulipas, San Luis Potosi and Veracruz, provided significant new distribution 
records or data for seven freshwater decapods species. Information on each 
species is presented, including restricted synonymy, previously known distri- 
bution, new localities, and notes on the significance of the new data. 



Villalobos-Hiriart et al. (1993) reported 
that Mexican freshwater decapods com- 
prised about 132 species. Of these, many 
are known only from the type locality and, 
in several cases, knowledge on habitat is 
scarce or non-existent. Rapid changes in 
habitat conditions, introduction of non-na- 
tive species, and decline in abundance of 
some elements of the original crustacean 
decapod fauna in northern Mexico (see 
Contreras-Balderas 1991; Rodriguez-Al- 
maraz & Campos 1994) prompted us to up- 
date the diversity, ranges, and habitat of the 
22 species reported from that region. Col- 
lections in selected freshwater habitats in 
the states of Nayarit, Nuevo Leon, Tamau- 
lipas, San Luis Potosi, and Veracruz pro- 
duced significant new distribution data for 
seven freshwater decapods species one 
Atyidae, one Cambaridae and five Palae- 
monidae). Abbreviations used are: UANL, 
Arthropoda Collection, Facultad de Cien- 
cias Biologicas, Universidad Autonoma de 
Nuevo Leon; UABC, Invertebrate Collec- 
tion, Facultad de Ciencias, Universidad Au- 
tonoma de Baja California; IBUNAM-EM, 
Crustacean Collection, Instituto de Biolo- 
gia, Universidad Nacional Autonoma de 
Mexico. For each species listed, a restricted 
synonymy is provided and includes original 



description or redescription, changes in ge- 
neric assignment, and known distribution. 

Family Atyidae 
Potimirim mexicana (De Saussure, 1857) 

Potimirim mexicana. — Villalobos-Figueroa, 
1959:295-313, pi. VI, figs. 29-38, pi. 
VII, figs. 39-50, pi. VIII, figs. 51-54, pi. 
IX, figs. 55, 56, map. 2, 3.-1982:217, 
223.— Hart, 1961:67, 76.— Smalley, 1963: 
177, 179, 181.— Villalobos-Hiriart et al., 
1993:281. 

Known distribution. — Mexico. This spe- 
cies is found near the sea in the hydrolog- 
ical system of the Gulf of Mexico slope, 
from Rio Soto la Marina basin, Tamaulipas, 
to Rio Grijalva basin. Tabasco, and rivers 
emptying in Laguna del Carmen, Campe- 
che. Central America. Honduras, Costa 
Rica. Antilles. Cuba, Jamaica and Puerto 
Rico (Villalobos-Figueroa 1982). 

Material examined. — 21 females, Estero 
de Canala, Guayabitos, Nayarit, 2r05'N, 
105°15'W, 1 Jun 1995 (UANL). 8 males, 13 
females, Camino a las Varas, Nayarit, 
21°10'N, 105°14'W, 2 Jun 1995. 4 males, 
Rio Coy (Rio Choy) [Rio Panuco basin], 28 
km south of Valles city, San Luis Potosi, 
2r51'N, 98°56'W, 27 Aug 1994 (UANL). 
3 males (topotypes, IBUNAM-EM uncat.). 



VOLUME 109, NUMBER 1 



35 



M. A. Quevedo Park, Playa Norte, Vera- 
cruz, Veracruz, 19°14'N, 96°08'W, 1958. 

Remarks. — Our specimens agree with 
Villalobos-Figueroa's (1959) description 
except that we found up to 15 uncinuli on 
the appendix interna and, like in P. poti- 
mirim (Miiller), a plumose seta close to it. 
Even so, morphometric and shape features 
recorded by Villalobos-Figueroa allow easy 
separation of these two species. This record 
represents the farthest inland finding of P. 
mexicana in the Gulf of Mexico slope, ap- 
proximately 250 km upstream in the Rio 
Panuco-Rio Coy basin, and is the first re- 
port of this species in the Mexican Pacific 
slope. 

Family Cambaridae 

Procambarus roberti 

Villalobos-Figueroa & Hobbs, 1974 

Procambarus (Pennides) roberti Villalo- 
bos-Figueroa & Hobbs, 1974:8, fig. 2.— 
Hobbs, 1989:74. 

Known distribution and habitat. — Ditch 
from La Media Luna, 4.8 miles (7.7 km) 
south of Rio Verde (on highway to Pedro 
Montoya) and 2.5 miles (4 km) west on dirt 
road to Mina El Refugio, San Luis Potosi 
(type locality). In streams. 

Material examined. — 1 male I, 3 fe- 
males, 6 juvenile males, 7 juvenile females. 
El Venado, San Luis Potosi (151 northwest 
of San Luis Potosi city and 105 km south- 
west of Matehuala city, San Luis Potosi), 
22°58'N, 101°04'W, Feb 1994 (UANL). 

Remarks. — The habitat of Procambarus 
roberti at El Venado is a spring-pool of ap- 
proximately 20 m^ with clear water and a 
rocky bottom. Only scarce grasses were ob- 
served in the margin of the pool amongst 
which the crayfishes were collected. Ostra- 
cods of the genus Darwinula and Cypron- 
odopsis were in the sediment of the bottle 
where the crayfishes were fixed; however, 
the entocytherid symbiont, Ankylocythere 
barbouri Villalobos-Figueroa & Hobbs, 
1974, was not found. It is not clear how P. 
roberti moved 160 km northwest from La 



Media Luna to El Venado since no super- 
ficial aquatic drainage system connects 
these localities. An hypogean dispersion or 
human introduction are two possible expla- 
nations. 

Family Palaemonidae 

Macrobrachium acanthurus 

(Wiegmann, 1836) 

Palaemon acanthurus Wiegmann, 1836: 
150. 

Macrobrachium acanthurus, Hedgpeth, 
1949:30, figs, la, 2, 5.— Holthuis, 1952: 
45-53, pi. 8, 9, figs, a, b.— Villalobos- 
Figueroa, 1982:217, 224.— Williams, 
1984:66-68, figs. 44, 45. — Markham et 
al., 1990:419. 

Known distribution and habitat. — From 
North Carolina, U.S.A. to Brazil (Holthuis 
1952; Williams 1984). In Mexico it ranges 
from southern Tamaulipas to Quintana Roo 
(Markham et al. 1990). The species lives in 
fresh or sometimes brackish water, and gen- 
erally is not found far inland (Holthuis 
1952). 

Material examined. — 3 males, Rio Ra- 
mos [San Juan basin], municipality of Al- 
lende, Nuevo Leon, 25°16'N, 100°00'W, 15 
Apr 1992 (UANL). 

Remarks. — This is the first record of M. 
acanthurus in the state of Nuevo Leon and 
represents the farthest inland finding of this 
species in Mexico, approximately 360 km 
upstream in the Rio Bravo-Rio San Juan ba- 
sin. 

Macrobrachium hobbsi 

Nates-Rodriguez & Villalobos-Hiriart, 

1990 

Macrobrachium hobbsi Nates-Rodriguez & 
Villalobos-Hiriart, 1990:7, Fig. 3.— Vil- 
lalobos-Hiriart et al., 1993:285. 

Know distribution. — Los Tuxtlas area, 
southern Veracruz; Arriaga and Tonala, 
Chiapas; Rio Ostuta, Oaxaca, and Rio Mur- 
ga near Petatlan, Guerrero; presumably to 



36 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Nayarit (Nates-Rodriguez & Villalobos- 
Hiriart 1990; Villalobos-Hiriart 1993). 

Material examined. — 4 males, 1 female, 
20 juveniles, Rio Coy ( = Rio Choy), 28 km 
south of Ciudad Valles, San Luis Potosi, 
Apr, Jun 1994 (UANL); 1 male, 1 female, 
Rio Huichihuayan, 70 km south of Ciudad 
Valles, San Luis Potosi, 2r32'N, 98°56'W, 
Apr 1994 (UANL); 7 juveniles, small pool, 
km 160 on the highway Tuxpam-Tampico, 
(30 km south of Panuco [Ozuluama]), Ve- 
racruz, 2r5rN, 97°48'W, Nov 1993 
(UANL); 3 juveniles, small pool, km 240 
on highway Poza Rica-Tuxpam, Veracruz, 
20°45'N, 97°31'W, Nov 1993 (UANL). 

Remarks. — The presence of M. hobbsi in 
northern Veracruz and San Luis Potosi rep- 
resents a northern range extension of ap- 
proximately 850 km. It is remarkable that 
juveniles of M. hobbsi were found in the 
same habitat with Palaemonetes mexicanus 
Strenth, at Rio Coy, San Luis Potosi. This 
does not agree with Strenth's (1976) con- 
clusion that juveniles of Macrobrachium 
competitively exclude Palaemonetes. 

Macrobrachium olfersi 
(Wiegmann, 1836) 

Palaemon olfersii Wiegmann, 1836:150. 

Macrobrachium olfersi. — Holthuis, 1952: 
95-103, pi. 24, pi. 25, figs, a, b.— Hedg- 
peth, 1949:35, figs. Id, 4, 5.— Villalobos- 
Figueroa, 1 967: 1 67-1 7 1 .—Villalobos- 
Hiriart, 1993:285. 

Known distribution. — Western Atlantic 
slope. Lower Cape Fear River near South- 
port, North Carolina; Florida; Louisiana; 
Texas; and southern Veracruz, Mexico to 
Santa Catarina, Brazil (Holthuis 1952). 
Eastern Pacific slope. From Chiapas to 
Nayarit, Mexico (Villalobos-Hiriart et al. 
1993). 

Material examined. — 1 male, 3 juveniles, 
Rio Limon, 8 km north of Ciudad Mante, 
Tamaulipas, 22°49'N, 98°56'W, 12 Jun 
1994 (UANL). 

Remarks. — The identification of M. ol- 
fersi was possible by comparing the adult 



male above noted with the description and 
figures provided by Holthius (1952). The 
shrimps were collected in a shallow water 
area (15 cm depth) of the Rio Limon, 
amongst aquatic vegetation. This records 
represent the farthest inland finding of this 
species in Mexico, approximately 205 km 
upstream in the Rio Guayalejo-Rio Limon 
basin. 

Palaemonetes kadiakensis Rathbun, 1902 

Palaemonetes kadiakensis Rathbun, 1902: 
93.— Strenth, 1976:2.-1994:91. 

Palaemonetes {Palaemonetes) kadiakensis, 
Holthuis, 1949:92.-1952:212, pi. 51, figs, 
k-n, pi. 52, figs, a, b. — Smalley, 1964: 
23 1 .— Flemming, 1969:444.— Villalobos- 
Figueroa & Hobbs, 1974:15, fig. 8b. 

Known distribution. — This species is 
widespread in fresh-water habitats of Cen- 
tral U.S.A., including the shores of lakes 
Ontario, Erie, and Michigan, the Mississip- 
pi basin, and the basin of several rivers 
emptying east and west of the Mississippi 
river in the Gulf of Mexico (Holthuis 
1952). The only Mexican record is that of 
Creaser (1932) for the state of Nuevo Leon, 
northeast Mexico (Strenth 1976). 

Material examined. — 4 males, 5 females 
(1 ovigerous), Rio Sabinas Hidalgo, Valle- 
cillos, Nuevo Leon, 26°39'N, 99°59'W, 20 
Apr 1979 (UANL). Tamaulipas: 21 males, 
15 females, 13 Mar 1981 (UABC); 1 male, 

3 ovigerous females, 22 Oct 1981; 1 male, 

4 ovigerous females, 22 Nov 1981; 1 male, 
6 females, 30 Jun 1982, Rio Alamo, Cd. 
Mier, 26°25'34"N, 99°06'41"W; 1 male, 4 
females (2 ovigerous), 23 Oct 1983, Presa 
Falcon, Miguel Aleman, 26°39'N, 99°12'W; 
1 male, 2 females, 30 Mar 1983; 1 male, 4 
females, 14 Mar 1987; Rio Bravo, Mata- 
moros 25°50'N, 97°24'W (UANL). 

Remarks. — Based on the present record 
we believe that the material reported by 
Creaser (1932) from Nuevo Leon came 
from the Rio Alamo-Rio Salado basin 
(northern Nuevo Leon), which empties into 
the Rio Bravo del Norte (Rio Grande) in 



VOLUME 109, NUMBER 1 



37 



the state of Tamaulipas. Previous collecting 
efforts in Central Nuevo Leon (Rio San 
Juan basin) failed to produce specimens of 
P. kadiakensis. 

The material of the Rio Alamo (13 Mar 
1981) was collected in shallow (<1 m 
depth) and clear-greenish water, the bottom 
was muddy and the aquatic vegetation in- 
cluded Chara sp. Additional information on 
the habitat is noted in Ruiz-Campos et al. 
(1985). 

Palaemonetes mexicanus Strenth, 1976 

Palaemonetes mexicanus Strenth, 1976:7, 
fig. 3.-1994:291. 

Known distribution and habitat. — Small 
spring-fed river, 15 miles (24 km) west of 
Ciudad Valles, San Luis Potosi (type local- 
ity). 

Material examined. — 7 males, 20 fe- 
males (5 ovigerous), Rio Coy, 28 km south 
of Ciudad Valles, San Luis-Potosi, Apr, Jun 
1994 (UANL). 

Remarks. — This is apparently the first 
record of P. mexicanus since its original de- 
scription by Strenth (1976). The new local- 
ity is about 50 km southeast of the type 
locality. Hobbs & Hobbs (1989) reported P. 
mexicanus to Mante river (Ciudad Mante, 
Tamaulipas); however, Strenth (1994) 
pointed out that Hobbs & Hobbs' speci- 
mens are of P. hobbsi Strenth, 1994. 

Five ovigerous females (cl. 11.4-18.5 
mm) were carrying 13 to 31 eggs having a 
diameter of 0.89-1.12 mm {X = 0.99) by 
1.15-1.4 mm (X = 1.25). 

Acknowledgments 

Financial support for a short-term visit to 
the Crustacean Collection, Instituto de Biol- 
ogia, Universidad Nacional Autonoma de 
Mexico (UNAM) was provided by Direc- 
cion de Intercambio Academico, UNAM to 
GAR-A. Our great appreciation is given to 
J. F Fitzpatrick Jr., H. H. Hobbs III, N. E. 
Strenth and J. L. Villalobos-Hiriart for their 
criticism and helpful comments on the 



manuscript. EC is a fellow of the "Progra- 
ma de Estimulo al personal Academico 94/ 
95" of the Universidad Autonoma de Baja 
California (UABC). This is contribution 
number 4 of the program "Crustaceos De- 
capodos (formerly Cambarinos) del Norte 
de Mexico" of the UABC and Universidad 
Autonoma de Nuevo Leon. 

Literature Cited 

Contreras-Balderas, S. 1991. Conservation of Mexi- 
can freshwater fishes: some protected sites and 
species and recent federal legislation. Pp. 191- 
197 in W. L. Minckley and J. E. Deacon, eds.. 
Battle against extinction: native fish manage- 
ment in the American West. University of Ari- 
zona Press, Tucson, Arizona. 

De Saussure, H. 1857. Diagnoses de quelques Crus- 
taces nouveaux de I'Amerique tropical. — Re- 
vue et Magasin de Zoologie pure et appliquee 
2(9):501-505. 

Creaser, E. P. 1932. The decapod crustaceans of Wis- 
consin. — Transactions of the Wisconsin Acad- 
emy of Sciences, Arts and Letters 27:321-338. 

Flemming, L. E. 1969. Use of male external genitalic 
details as taxonomic characters in some species 
of Palaemonetes (Decapoda, Palaemonidae). — 
Proceedings of the Biological Society of Wash- 
ington 82:443^52. 

Hart, C. W. 1961. The freshwater shrimps (Atyidae 
and Palaemonidae) of Jamaica, W.L with a dis- 
cussion of their relation to the ancient geogra- 
phy of the Western Caribbean area. — Proceed- 
ings of the Academy of Natural Sciences of 
Philadelphia 113(4):61-80. 

Hedgpeth, J. W. 1949. The North American species 
of Machrobrachium (River shrimps). — Texas 
Journal of Science 1:28-38. 

Hobbs, H. H. Jr. 1989. An illustrated checklist of the 
American crayfishes (Decapoda: Astacidae, 
Cambaridae, and Parastacidae). — Smithsonian 
Contribution to Zoology 480:1-235. 

, & H. H. Hobbs III. 1989. New locality re- 
cords for two poorly known mexican freshwater 
shrimps (Decapoda, Palaeomonidae). — Crusta- 
ceana 57(2):220-222. 

Holthuis, L. B. 1949. Note on the Palaemonetes 
(Crustacea Decapoda) found in the United 
States-of America. — Proceedings of the Kon- 
inklinjke Nederlandse Akademie van Weten- 
schappen 52:87-95. 

. 1952. The subfamily Palaemoninae. A gen- 
eral revision of the Palaemonidae (Crustacea 
Decapoda Natantia) of the America, IL — Allan 
Hancock Foundation Occasional Papers 12:1- 
396. 



38 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



Markham, J. C, F. E. Donath-Hernandez, J. L. Villa- 
lobos-Hiriart, & A. Cantu Diaz-Barriga. 1990. 
Notes on the shallow-water marine Crustacea of 
the Caribbean coast of Quintana Roo, Mexi- 
co. — Anales del Instituto de Biologia, Univer- 
sidad Nacional Autonoma de Mexico, Serie 
Zoologia 61(3):405^46. 

Nates-Rodriguez, J. C, & J. L. Villalobos-Hiriart. 
1990. Dos nuevas especies de camarones de 
agua dulce del genero Macrobrachium Bate, 
(Crustacea, Decapoda, Palaemonidae), de la 
vertiente occidental del Pacifico. — Anales del 
Instituto de Biologia, Universidad Nacional Au- 
tonoma de Mexico, Serie Zoologia, 61(1):1-11. 

Rathbun, M. J. 1902. Description of new Decapod 
Crustaceans from the West coast of North 
America. — Proceeding of the United States Na- 
tional Museum 24:885-905. 

Rodriguez- Almaraz G. A., & E. Campos. 1994. Dis- 
tribution and status of the crayfishes (Cambar- 
idae) of Nuevo Leon, Mexico. — Journal of 
Crustacean Biology 14(4):729-735. 

Ruiz-Campos, G., M. Torres-Morales and S. Contrer- 
as-Balderas. 1985. Peces del Rio Alamo, Sub- 
cuenca del Bravo, Mexico. II. Estructura y din- 
amica de la comunidad ictica. — Instituto de In- 
vestigaciones Cientificas, Universidad Autono- 
ma de Nuevo Leon (Mexico) 2:51-75. 

Smalley, A. E. 1963. The genus Potimirim in Central 
America (Crustacea, Atyidae). — Revista de 
Biologia Tropical 11(2):177-183. 

. 1964. A new Palaemonetes from Mexico 

(Decapoda: Palaemonidae). — Crustaceana 6: 
229-232. 

Strenth, N. E. 1976. A review of the systematics and 
zoogeography of the freshwater species of Pa- 
laemonetes Heller of North America (Crustacea: 
Decapoda). — Smithsonian Contribution to Zo- 
ology 228:1-28. 



. 1994. A new species of /'a/aemonerei' (Crus- 
tacea: Decapoda: Palaemonidae) from north- 
eastern Mexico. — Proceedings of the Biological 
Society of Washington 107:291-295. 

Villalobos-Figueroa, A. 1959. Contribucion al cono- 
cimiento de los Atyidae de Mexico. II. (Crus- 
tacea, Decapoda). Estudio de algunas especies 
del genero Potimirim (Ortmannia), con descrip- 
cion de una especie nueva en Brasil. — Anales 
del Instituto de Biologia, Universidad Nacional 
Autonoma de Mexico 30:269-330. 

. 1967. Estudio de los Palaemonidos de Mex- 
ico I. Macrobrachium acanthochirus una nueva 
especie del SE de Mexico. — Anales del Insti- 
tuto-de Biologia, Universidad Nacional Auton- 
oma de Mexico, Serie Zoologia, 26(1/2): 167- 
174. 

. 1982. Decapoda. Pp. 215-239 in S. H. Hurl- 

bert and A. Villalobos-Figueroa, eds.. Aquatic 
biota of Mexico, Central America and the West 
Indies. Aquatic Biota-San Diego State Univer- 
sity Foundation, San Diego California. 

, & H. H. Hobbs, Jr. 1974. Three new crusta- 



ceans from La Media Luna, San Luis Potosi, 
Mexico. — Smithsonian Contribution to Zoology 
174:1-18. 

Villalobos-Hiriart, J. L., A. Cantu Diaz-Barriga, & E. 
Lira-Fernandez. 1993. Los Crustaceos de Agua 
Dulce de Mexico. — Revista de la Sociedad 
Mexicana de Historia Natural 44:267-290. 

Wiegmann, A. F A. 1836. Beschreibung einiger 
neuen Crustaceen des Berliner Museums aus 
Mexiko und Brasilien. — Archiv fiir Naturges- 
chichte 2(1): 145-151. 

Williams, A. B. 1984. Shrimps, lobsters and crabs of 
the Atlantic coast of the eastern United States, 
Maine to Florida. Smithsonian Institution Press, 
Washington D.C., 550 pp. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(l):39-43. 1996 

Neocrangon zacae (Chace, 1937) synonymized with A^. resima 

(Rathbun, 1902), and compared with A^. communis (Rathbun, 1899) 

(Decapoda: Caridea: Crangonidae) 

Mary K. Wicksten 

Department of Biology, Texas A&M University, 
College Station, Texas 77843, U.S.A. 

Abstract. — Features previously used to distinguish between Neocrangon za- 
cae and A^. resima overlap greatly. The rostrum can be low and narrow, slightly 
elevated and deep, or high with a plate with or without teeth. The width of the 
hand of the first pereopod also varies, as does the distal width of the antennal 
scale. Similar variability exists in the shape of the rostrum of A^. communis, 
but this species has a prominent carina on the fifth abdominal somite and has 
broader hands of the first pereopod and a more horizontal dactyl. Usually, adult 
A^. resima are smaller than A^. communis. Neocrangon zacae is considered to 
be a junior synonym of N. resima. A redescription of Neocrangon resima is 
presented. 



Crangonid shrimp of the genus Neocran- 
gon Zarenkov, 1965, are characterized as 
follows: two median carapace spines, with- 
out submedian spines between the middor- 
sal line and hepatic spine, the branchioste- 
gal spine not very strong or flared sideways, 
the gastric region not depressed below the 
general level of the carapace, one or two 
spines at the distal end of the merus of first 
pereopod, second pereopods subchelate and 
subequal in length to other pereopods, 
fourth and fifth pereopods with normal dac- 
tyls; not broadened; lateral surface of the 
abdomen not sculptured. Four species have 
been reported from the northeastern Pacific: 
A^. communis (Rathbun, 1899), N. resima 
(Rathbun, 1902), N. abyssorum (Rathbun, 
1902) and N. zacae (Chace, 1937). Of 
these, A^. abyssorum is the most distinct, 
having enlarged eyes, a bent antennular pe- 
duncle, a long and narrow rostrum and a 
thin exoskeleton. It generally occurs on the 
lower continental slope, but has been re- 
ported as shallow as 97 m (Butler 1980). 

Colleagues conducting environmental 
impact surveys had difficulty distinguishing 



between N. communis, N. resima and A^. za- 
cae. Neocrangon resima has been recog- 
nized by its elevated rostrum with a ventral 
plate (Rathbun 1904:fig. 65). However, 
Rathbun noted that the development of the 
rostrum seemed to be dependent on age, 
with specimens smaller than 20 mm not 
showing this feature. Schmitt (1921) quoted 
Rathbun as stating that the rostrum of A^. 
communis also could be variable, with some 
specimens regenerating a narrow rostrum 
with an elevated tip. The illustration of the 
rostrum given by Schmitt (1921 :fig. 64) 
shows a bilobed rostrum with a small tooth, 
while that by Green & Butler (1988) shows 
a rostrum ending in a vertical plate. The 
rostrum of A^. zacae was described as as- 
cending at a slight angle "which varies 
somewhat" (Chace 1937). 

The hands of the first pereopod of N. re- 
sima were reported by Schmitt (1921) to be 
shorter and with a more longitudinal margin 
than in A^. communis. The hands of the first 
pereopod of A^. zacae were said by Chace 
(1937) to have a "longer palm" than those 
of N. resima, as well as a more longitudinal 



40 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



dactyl. However, Green & Butler (1988) did 
not indicate any difference between the pro- 
portions of the palm of their specimen of 
A^. resima and that of any other species. 

Schmitt quoted Rathbun as saying that N. 
resima had a "slight" median carina on the 
fifth abdominal somite while N. communis 
had a "blunt" carina. Green & Butler 
(1988) illustrated a specimen of A^. resima 
with a "moderate" median dorsal carina, 
but noted that the carapace and abdominal 
sulci bore a short pubescence, as Butler 
(1980) reported for N. communis. Neocran- 
gon zacae lacked a carina on the fifth ab- 
dominal somite (Chace 1937). 

The ranges and habitats of the three spe- 
cies have been reported to overlap in the 
eastern Pacific. According to Butler (1980), 
N. communis ranges from the Bering Sea to 
San Diego, California at 16-1537 m on 
mud. Green & Butler (1988) reported N. re- 
sima from mud or mixed mud and sand bot- 
toms from Alice Arm, British Columbia to 
San Domingo Point, Baja California; 
Schmitt (1921) reported the depth range as 
28-487 m. Neocrangon zacae has been re- 
ported from Monterey Bay, California to 
north of Gorgona Island, Colombia (al- 
though most reports come from California, 
U.S.A. and Baja California, Mexico) at 
depths from shore to 572 m. It has been 
reported from muddy or rocky bottoms. In 
southern California and Baja California, N. 
zacae usually was collected at shallower 
depths than N. communis (Wicksten 1980). 
Neocrangon communis and N. resima have 
been reported to be collected at the same 
trawling station (Schmitt 1921; Green & 
Butler 1988). 

Considering the overlap in range, depth 
and morphological features reported for A^. 
communis, N. zacae and N. resima. I ex- 
amined specimens to determine what con- 
sistent differences could be found between 
them. Comparison of these specimens in- 
dicate that only two of these species are dis- 
tinct. 



Materials and Methods 

I examined 179 specimens identified as 
A^. communis, N. resima or N. zacae from 
Santa Monica Bay and Monterey Bay, Cal- 
ifornia and Cook Inlet and the northeastern 
Gulf of Alaska. One hundred fifty two spec- 
imens from Santa Monica Bay were loaned 
by Ann Dalkey of the Hyperion Treatment 
Plant, Los Angeles County Environmental 
Monitoring Division; the specimens from 
Monterey Bay and Alaska were loaned by 
the California Academy of Sciences (CAS 
014038, 020155, 020163 and 020155). I 
also examined the six syntypes of N. resi- 
ma, collected off San Diego, California, 
from the National Museum of Natural His- 
tory, Smithsonian Institution (USNM 
25246). 

For each specimen, I noted total length, 
shape (low, deep, with plate, with or with- 
out teeth) and elevation of the rostrum, 
presence or absence of a noticeable dorsal 
carina of the fifth abdominal somite, and 
shape of the hand of the first pereopod. 
These data were tabulated and compared 
with the original descriptions of the species. 

Results 

The most striking difference among the 
shrimp examined was that all specimens 
from the Gulf of Alaska and Cook Inlet had 
a pronounced and blunt dorsal carina on the 
dorsal surface of the fifth abdominal somite 
(Fig. If). None of those from California had 
more than a vague trace of a dorsal carina 
(Fig. Ic). The Alaskan specimens were 
larger, ranging from 37.1-61.1 mm in total 
length. The specimens from California 
ranged from 19.5-50.6 mm in total length, 
but most individuals (59) were between 35- 
40 mm long. 

The shape of the rostrum was low and 
narrow in 87 specimens from California 
and 13 Alaskan specimens, elevated but 
without a plate in 52 specimens from Cal- 
ifornia and eight from Alaska, and elevated 
and with a plate in 14 specimens from 
Santa Monica Bay, three from Monterey 



VOLUME 109, NUMBER 1 



41 


















f 



Fig. 1. Neocrangon resima (Rathbun, 1902). a, 10 shapes of the rostrum; b, subchela of first pereopod; c, 
dorsal surface of fifth abdominal somite in frontal view. Neocrangon communis, d, three shapes of the rostrum; 
e, subchela of first pereopod; f, dorsal surface of fifth abdominal somite in frontal view. 



42 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Bay and one from Alaska. In other speci- 
mens, the rostrum was broken off. In the 
collection from Santa Monica Bay, speci- 
mens with an elevated rostrum with a plate 
comprised about 9% of the total collection. 

Some of the shapes of the rostrum are 
shown in Fig. la and Id. It was difficult to 
determine the difference between "elevated 
and deep" and "elevated with a plate"; 
moreover, the number of teeth (if any) also 
varied. Most of the rostra did not have teeth 
except for a pointed apex. Among the syn- 
types of A^. resima, the shape of the rostrum 
varied: two had a very low rostrum, one 
had a short but upraised rostrum and three 
had a moderately upraised rostrum. It is 
noteworthy that none of the syntypes had a 
raised rostrum with a plate, as illustrated by 
Rathbun (1904:fig. 65b). 

Most of the specimens from Alaska had 
a broader hand and also a more horizontal 
dactyl of the first pereopod than those from 
California (Fig. le). In general, the speci- 
mens from California had a more elongated 
hand with a more vertical dactyl (Fig. lb), 
similar to that illustrated by Chace (1937: 
fig. 9d) for N. zacae. 

Discussion 

In comparing the syntypes of A^. resima, 
the other specimens and the descriptions, 
the presence or absence of a carina on the 
dorsal surface of the fifth abdominal somite 
was the most reliable distinguishing feature 
for recognition of species. Such a carina is 
present in A^. communis. Neocrangon com- 
munis also may have a broader hand of the 
first pereopod, about 2.7-3 X long as wide, 
although some specimens may have a hand 
as much as 3.7 X long as wide (Butler 
1980). The hand has a dactyl that closes 
somewhat horizontally. The specimens 
from Alaska could be confidently identified 
as N. communis. The specimen illustrated 
by Green & Butler (1988) as N. resima 
probably also belongs to N. communis. Ex- 
cept for the variation in the shape of the 
rostrum, the specimens fit well the descrip- 



tion of A^. communis given by Butler (1980, 
as C rang on communis). 

The shape of the rostrum is unreliable as 
a diagnostic character of species of Neo- 
crangon. Among the specimens from Mon- 
terey Bay and Santa Monica Bay, it varied 
from low to high, shallow to deep, and with 
or without a plate, which sometimes had 
teeth. Variation also occurred among the 
syntypes of N. resima and the specimens of 
A^. communis. 

Rathbun (1904) stated that the develop- 
ment of the rostral plate was dependent on 
age, "specimens 20 mm long show no ev- 
idence of it." However, in two specimens 
with a total length of 22-28 mm, there was 
an elevated rostrum; another had a rostrum 
with a well-developed plate. Because my 
samples contained only one specimen less 
than 20 mm long, it is difficult to say 
whether or not the development is related 
to age or size. The smallest specimen, 19.5 
mm long, had a low rostrum without a 
plate. 

The features of the fifth abdominal so- 
mite and first pereopod are indistinguish- 
able between A^. resima and N. zacae. The 
ranges as reported for the two species, their 
habitat and depth also overlap. In the ab- 
sence of any other distinguishing features, 
Neocrangon zacae (Chace, 1937) must be 
considered to be a junior synonym of N. 
resima (Rathbun, 1902). It is unfortunate 
that the species was illustrated in Rathbun's 
work from an atypical specimen having a 
raised rostrum and a ventral plate, for this 
shape of the rostrum occurs in less than 
10% of the population. A redescription of 
N. resima, including new information on 
variability within the species, follows. 

Neocrangon resima (Rathbun, 1902) 

Crangon resima Rathbun, 1902: 889. — 

Rathbun 1904:124, fig. 65. 
Crago resima. — Schmitt 1921:96, fig. 64. 
Crago zacae Chace, 1937:136, text-fig. 9. 
Neocrangon zacae. — Kuris & Carlton 

1977:554.— Wicksten 1980:39. 



VOLUME 109, NUMBER 1 



43 



Neocrangon resima. — Kuris & Carlton 
1977:554. 

Redescription. (Modified from Chace 
1937). — Rostrum usually narrow, with pro- 
nounced dorsal sulcus and blunt apex, vari- 
able in length, not reaching end of cornea 
of eye to exceeding entire eye. Rostrum as- 
cending, usually at slight angle but as much 
as 45° in some specimens; ventral plate, 
with or without teeth, may be present. 

Carapace about 0.25 X length of abdo- 
men. Gastric region not depressed below 
general outline of carapace. Two spines in 
dorsal midline, posterior one longer and 
arising in front of middle of carapace. 
Strong hepatic spine in line with anterior 
dorsal spine. Anterior margin with antennal 
and branchiostegal spines. 

Eye large and black. 

Abdominal somites with few small setae 
along ventral surface but without obvious 
pubescence. Somites 1 and 2 with elevated 
posterior portions, elevations preceded by 
broad, shallow transverse sulci. Somites 1- 
4 without carinae. Pleura of first to third 
somites with rounded or squared margins, 
fourth pleuron with minute posteroventral 
spine, fifth pleuron with posterolateral 
point. Fifth somite occasionally with ob- 
scure median carina. Sixth somite approxi- 
mately 1.6X length of fifth, bearing pair 
dorsal longitudinal carinae separated by 
median sulcus and bounded by shallower 
lateral sulci. Telson longer than sixth so- 
mite, with slight median sulcus, bearing 2 
pairs dorso-lateral spines, apex blunt. 

Antennular peduncle with 3 segments, 
second 2-3 X length of third, stylocerite 
short. Antennular flagella exceeding anten- 
nal scale. Basicerite of antenna with lateral 
spine. Spine of antennal scale exceeding 
blade, antennal scale about 3.5 X long as 
wide. 

Third maxilliped long and slender, ex- 
tending slightly beyond antennular flagella, 
exopod with lash. First pereopod subche- 
late, merus with spine at outer angle of dis- 
tal end, palm of subchela 3-3.3 X long as 



wide, dactyl closing obliquely. Second per- 
eopods slender, shorter than first, chelate. 
Third pereopods slender, extending well be- 
yond third maxillipeds, fourth and fifth per- 
eopods shorter; all with simple, slender dac- 
tyls. Uropods equal to or exceeding length 
of telson. 

Color in life translucent, mottled finely 
with greenish-brown and scarlet on dorsal 
and lateral surfaces; ventral surface white, 
antennae banded with scarlet and white, 
uropods and telson mottled. 

Literature Cited 

Butler, T. H. 1980. Shrimps of the Pacific coast of 
Canada. — Canadian Bulletin of Fisheries and 
Aquatic Sciences 202:1-280. 

Chace, F. A. Jr. 1937. The Templeton Crocker Expe- 
dition. VII. Caridean decapod Crustacea from 
the Gulf of California and the west coast of 
Lower California. — Zoologica 22:109-138. 

Green, G., & T. H. Butler. 1988. Range extensions of 
three caridean shrimps to British Columbia wa- 
ters. — Royal British Columbia Museum Contri- 
butions to Natural Science 8:1-7. 

Kuris, A. M., & J. T. Carlton. 1977. Description of a 
new species, Crangon handi, and new genus, 
Lissocrangon, of crangonid shrimps (Crustacea: 
Caridea) from the California coast, with notes 
on adaptation in body shape and coloration. — 
Biological Bulletin 153:540-559. 

Rathbun, M. J. 1 899. List of Crustacea known to oc- 
cur on and near the Pribilof Islands. Pp. 555- 
557 in D.S. Jordan et al., ed.. The fur seals and 
fur seal islands of the north Pacific Ocean, 
Washington, D.C., Pt. 3. 

. 1902. Descriptions of new decapod crusta- 
ceans from the west coast of North America — 
Proceedings of the U.S. National Museum 24: 
885-905. 

. 1904. Decapod crustaceans of the northwest 



coast of North America. — Harriman Alaska Ex- 
pedition, Series 10:1—210. 

Schmitt, W. L. 1921. The marine decapod Crustacea 
of California. — University of California Publi- 
cations in Zoology 23:1^70. 

Wicksten, M. K. 1980. Range extensions of four spe- 
cies of crangonid shrimps in the eastern Pacific 
Ocean (Decapoda: Crangonidae). — Bulletin of 
the Southern California Academy of Sciences 
79:38-41. 

Zarenkov, N. A. 1965. Revision of the genera Cran- 
gon Fabricius and Sclerocrangon G.O. Sars 
(Decapoda, Crustacea). — Zoologicheskii Zhur- 
nal 44:1761-1775. (In Russian.) 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(l):44-52. 1996 

Daldorfia Rathbun, 1904 (Crustacea: Decapoda) from the 

Neogene of Japan 

Hiroaki Karasawa and Hisayoshi Kato 

(H. Karasawa) Mizunami Fossil Museum, Yamanouchi, Akeyo, Mizunami, Gifu 509-61, Japan; 
(H. Kato) Natural History Museum and Institute, Chiba, 955-2 Aobacho, Chuoku, 

Chiba 260, Japan 

Abstract. — Daldorfia nagashimai, new species, is described from the Hi- 
gashimorogata Formation (Upper Miocene - Lower Pliocene), Miyazaki Group 
of Miyazaki Prefecture, Japan. This species is characterized by a large carapace 
with irregular, large, granulose tubercles dorsally and without a deep, eroded 
area behind a longitudinally hexagonal mesogastric lobe. Another specimen 
representing a second unnamed species is recorded from the Aoso Formation 
(Upper Miocene) of Miyagi Prefecture, Japan. These species represent the first 
records of Neogene decapod crustaceans from Japan and extend the geologic 
range of the genus Daldorfia to the Late Miocene age. 



The subfamily Parthenopinae is a small 
group including eight Recent genera, Daira 
de Haan, 1833, Dairoides Stebbing, 1920, 
Daldorfia Rathbun, 1904, Leiolambrus A. 
Milne Edwards, 1878, Parthenope Weber, 
1795, Solenolambrus Stimpson, 1871, Thy- 
rolambrus Rathbun, 1894, and Tutankha- 
men Rathbun, 1925. The fossil record of 
Parthenope Weber, 1795, is robust in Ce- 
nozoic deposits throughout the world 
(Glaessner 1969). Daira is recorded from 
the Eocene-Miocene of Europe, the Plio- 
cene of Fiji, and the Miocene of Japan 
(Glaessner 1969; Muller 1984; Muller & 
Collins 1991; Karasawa 1993). The only 
known fossil Leiolambrus was described 
from the Upper Eocene Bartonian of En- 
gland (Quayle & Collins 1981) and Tut- 
ankhamen also has been reported from the 
Oligocene Gambier Limestone of Australia 
(Jenkins 1985). The fossil records of the 
other four Recent genera are unknown 
throughout the world, but the extinct, Me- 
solambrus declinatus Muller & Collins, 
1991, similar to Thyrolambrus Rathbun, 
1894, was described from the Szepvolgy 
Formation (Upper Eocene) of Hungary; 
Glaessner (1969) recognised the Upper Eo- 



cene, Phrynolambrus Bittner, 1893 as the 
junior synonym of the subgenus Pseudo- 
lambrus Paulson, 1875, in Parthenope, but 
subsequently Guinot (1979) separated 
Phrynolambrus from Pseudolambrus and 
suggested that Phrynolambrus has a close 
affinity with Daira and Dairoides. 

The purpose of this paper is to describe 
a new species and a related, but unnamed 
species of Daldorfia from the Neogene de- 
posits of Japan. The materials were collect- 
ed from a road cut (Loc. MYZ-4 of Kara- 
sawa 1993, 31°56'48"N, 131°16'46"E) at 
Akatani, Uranona, Takaoka-cho, Higashi- 
morogata-gun, Miyazaki Prefecture (Fig. 
lA). Siltstone of the Ay a Member of the 
Higashimorogata Formation, Miyazaki 
Group (Upper Miocene to Upper Pleisto- 
cene) is exposed at this locality (Tomida 
1991; Karasawa 1993). This formation is 
assigned to Zones N.17b-18 (latest Mio- 
cene-earliest Pliocene) of Blow's scale of 
planktonic foraminifera (Suzuki 1987). 
Daldorfia nagashimai, new species, is 
based upon the holotype and two paratype 
specimens included within calcareous nod- 
ules. Karasawa (1993) reported two species 
of crabs from this locality. The decapod as- 



VOLUME 109, NUMBER 1 



45 



Tohoku exp.' 




T 



Sinkansen Line 

/ / Yoshidagawa River 



w Fossil-bearing 
^ locality 



5 km 



I -^^-n O Sendai City- 





A. MIYAZAKI AREA 



O Miyazaki City 

'I 



The Sea of 
Hyuga 



Fig. 1. Map of Japan showing the fossil-bearing localities. 



semblage is dominated by Carcinoplax 
prisca Imaizumi. Two specimens of Linu- 
parus sp. aff. L. trigonus (von Siebold) 
have been found also. Tomida (1991) re- 



ported some molluscs, Perotrochus sp., 
Bathybembix sp., Hindsia sp. and Acila 
submirabilis Makiyama from the locality. 
Decapods collected from the locality sug- 



46 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



gest an environment within the lower sub- 
Httoral zone on a muddy to sandy bottom 
(Karasawa 1993). 

A large manus of the second species, 
Daldorfia sp., was obtained from the Aoso 
Formation at the exposed road-cut south of 
Matsugai, Tomiya Town, Miyagi Prefecture 
(38°20'22"N, 140°55'56"E) (Fig. IB). The 
Aoso Formation consists mainly of cross- 
laminated, poorly consolidated coarse- 
grained sandstone with remarkable key 
beds of tuff layer (K7) intercalated within 
the lower part and a pumiceous tuff layer 
underlying the conglomerate bed (K8) in 
the middle part of the Formation (Kitamura 
et al. 1983). The decapod fossil was ob- 
tained from this lithology beneath the K8 
beds. Fujiwara (1992) discussed the Late 
Miocene molluscan assemblages of the re- 
gion. He discriminated the Serripes-Miya 
and Glycymeris-Dosinia assemblages with- 
in the K8 beds, and regarded the inhabitants 
to be subneritic to mesoneritic, respectively. 
According to Fujiwara (1992) and Saito & 
Fujiwara (1994), the middle to the upper 
part of the Aoso Formation lying several 
meters above the K7 tuff bed is correlated 
to Blow's N17b Zone. Therefore, the geo- 
logic age of the fossil bearing horizon is 
assignable to the latest Miocene. From an- 
other locality of the Aoso Formation, a 
small number of fingers referable to the 
families Callianassidae and Parthenopidae 
were collected by the same person who dis- 
covered the material described below. 

Section Heterotremata Guinot, 1977 
Superfamily Parthenopoidea MacLeay, 

1838 

Family Parthenopidae MacLeay, 1838 

Subfamily Parthenopinae MacLeay, 1838 

Genus Daldorfia Rathbun, 1904 

Type species. — Cancer horridus Linnae- 
us, 1758, by monotypy (ICZN Opinion 
1582); Recent, Indo-West Pacific. 

Geologic range. — Late Miocene — Re- 
cent. 



Daldorfia nagashimai, new species 
Figs. 2-A 

Material.— KMNH IvP 300,022, holo- 
type (Kitakyushu Museum and Institute of 
Natural History, 6, Nishihonmachi 3-cho- 
me, Yahatahigashiku, Kitakyushu, 805 Ja- 
pan), coll. H. Nagashima, 1994; MFM83058, 
83059, 2 paratypes (Mizunami Fossil Mu- 
seum, Yamanouchi, Akeyo, Mizunami, 
Gifu, 509-61 Japan), coll. S. Tomida, 1993. 

Diagnosis. — Large parthenopid; dorsal 
carapace covered with irregular, large gran- 
ulose tubercles; mesogastric lobe large, lon- 
gitudinally hexagonal, without deep, eroded 
area behind it; major cheliped like Daldor- 
fia horrida, large, long, with stout fingers. 

Description. — A large Daldorfia; cara- 
pace appears to be broadly pentagonal in 
outline, width approximately 1.4 times 
length. Front broadly triangular, down- 
turned, with shallow, ovate dorsal hollow. 
Orbits small, subovate, directed anterolat- 
erally. Anterolateral margin strongly con- 
vex, bearing irregular, large tubercles; deep 
notch defined lateral termination of from 
cervical groove. Posterolateral margin 
strongly convex. Posterior margin obscured 
by poor preservation. 

Dorsal surface with inflated, tuberculate 
regions separated by shallow, moderately 
well-defined grooves. Protogastric lobes 
strongly convex, with sharply pointed tu- 
bercles on highest parts, deep depressions 
between gastric and hepatic lobes. Meso- 
gastric lobe with large, granulose tubercles, 
longitudinally hexagonal, gently convex; 
narrow anterior mesogastric process with 
tubercles longitudinally arranged; deep de- 
pression on either side of mesogastric lobe. 
Cardiac lobe broken. Highest part of strong- 
ly convex hepatic regions with sharply 
pointed tubercles. Branchial lobes uneven, 
with irregular, large, granulose tubercles 
and small setal pits. Intestinal region de- 
pressed, with irregular tubercles. 

Dactylus and propodus of right major 
cheliped (paratype) preserved, but tips of 
both fingers, and proximal end and mesial 



VOLUME 109, NUMBER 1 



47 




Fig. 2. Daldorfia nagashimai, new species, holotype, Xl.O; dorsal view; showing central and left side of 
carapace, parts of merus and carpus of left cheliped. 



surface of propodus broken. Fingers with a 
wide gape. Dactylus stout, with irregular, 
granulose tubercles; occlusal surface miss- 
ing. Imperfect fixed finger with a broad, 
flattened, molariform tooth on occlusal sur- 
face. Palm long, covered with longitudinal 
rows of irregular, conical, granulose tuber- 
cles decreasing in size proximally; greatest 
distal width about 1.5 times proximal 
width; tubercles large on dorsal and lateral 
surfaces, rather small on ventrolateral sur- 
face; ventral margin with conical, granulose 
tubercles; four longitudinal rows of spines 
present on ventromesial surface; two bro- 
ken bases of large mesial spines present, 
others missing. Carpus and merus of left 



cheliped (holotype) with granulose tuber- 
cles, merus bearing acute spines on dorsal 
margin. 

Pereiopods and ventral aspects unknown. 

Derivation of name. — The species name 
honors H. Nagashima, who collected the 
holotype. 

Remarks. — Daldorfia comprises nine Re- 
cent species from the Indo-Pacific and East 
Atlantic Oceans. Of these, Daldorfia na- 
gashimai most closely resembles D. horri- 
da (Linnaeus 1758) and D. rathbuni (de 
Man 1902), from the Indo-West Pacific 
Oceans, but the new species has a moder- 
ately uneven, not eroded dorsal surface with 
irregular, granulose tubercles. This new 



48 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Daldorfia nagashimai, new species, paratype, Xl.O. Dorsal view of carapace. 



species has a large, longitudinally hexago- 
nal mesogastric lobe with large, granulose 
tubercles, but D. horrida and D. rathbuni 
each have a small, rounded mesogastric 
lobe. Absence of a deep, eroded area behind 
the mesogastric lobe in the new species dis- 
tinguishes it from D. horrida and D. rath- 
buni. There is, in the general features of the 
major cheliped, considerable similarity be- 
tween D. nagashimai, and D. horrida: the 
latter bears two large, conical, distal tuber- 
cles, one on the lateral surface and the other 
on the dorsal surface of the propodus, but 
in D. nagashimai, the distal ends of the lat- 
eral and dorsal surfaces are covered with 
irregular, conical tubercles. Absence of 
spines on the mesial surface of the dactylus 
readily distinguishes D. nagashimai from 
D. horrida. 

Recent D. horrida and D. rathbuni differ 
from most other parthenopine crabs, includ- 
ing the other species of Daldorfia (i.e., D. 
investigatoris (Alcock 1895), D. spinosis- 
sima (A. Milne Edwards 1862)), in having 
both fingers of the major cheliped with 
large blunt molariform denticles on the 
proximal occlusal surfaces. 

Zipser & Vermeij (1978) observed that 
the Recent Daldorfia horrida uses the occlu- 
sal surfaces with molariform teeth of the ma- 



jor cheliped to crush gastropod shells (i.e., 
Cerithium, Cypraea, Drupa, Cantharus, 
Vasum) and to feed on hermit crabs. Ng & 
Rodriguez (1986) described detail the den- 
tition on both chelipeds of D. horrida. There 
is, on the occlusal surface of the major chela, 
a great similarity between the fossil species 
and Recent D. horrida. Thus, D. nagashimai 
appears to have acquired the crushing be- 
havior of feeding on gastropods and hermit 
crabs at least by the Late Miocene. 

Daldorfia sp. 
Fig. 5 

Material— SSME 13320 (Sendai Sci- 
ence Museum, 4-1, Forest Park, Dainohara, 
Aobaku, Sendai 981). 1 specimen, coll. Y. 
Takaizumi, 1991. 

Description. — A large propodus of left 
minor cheliped, oval in cross section, cov- 
ered on every surface except for fixed fin- 
ger with large, conical tubercles. Tubercles 
vary in size, up to 1 cm in diameter. Lateral 
surface has flattened tubercles, dorsal bor- 
der bears rather pointed ones. A hook- 
shaped projection with which carpus artic- 
ulates on dorsal border situated near prox- 
imal articulation. Fixed finger short, about 
'/s of total length, slightly curved inward. 



VOLUME 109, NUMBER 1 



49 



^• 



^-— ^,.Ji^:-- 




%^ 




Fig. 4. Daldorfia nagashimai, new species, paratype, X0.93. Right major cheliped: a, dorsal; b, lateral; c, 
mesial; d, ventral view. 



50 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 5. Daldorfia sp., X0.9. Propodus of left major cheliped: a, dorsal; b, lateral; c, mesial view. 



Stout with apex faintly curved downward. 
Occlusal surface with 3 blunt, subconical 
teeth. A shallow, short longitudinal furrow 
on apex below occlusal edge. 

Remarks. — In general shape, the present 
chela most closely resembles the minor 
chela of D. horrida, but differs in lacking 
granulose tubercles on the lateral surface 



and spines on the ventral margin. The mi- 
nor chela of D. nagashimai being as yet 
unknown, precludes comparison with D. sp. 

Acknowledgments 

We thank R. M. Feldmann (Kent State 
University) and J. S. H. Collins (London) 



VOLUME 109, NUMBER 1 



51 



for critically reading our manuscript, and H. 
Nagashima (Fukuoka City, Fukuoka), S. 
Tomida (Chukyo Gakuin University, Gifu), 
Y. Takaizumi (Sendai City, Miyagi) for of- 
fering their fossil decapod specimens. We 
also thank O. Fujiwara (Tono Geoscience 
Center, Gifu), Y. Okazaki (Kitakyushu Mu- 
seum and Institute of Natural History, Fu- 
kuoka), T. Komai (Natural History Museum 
and Institute, Cheba), and Y. Okumura (Mi- 
zunami Fossil Museum, Gifu), who offered 
much valuable advice. 

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Kitamura, N., A. Ozawa, & H. Nakagawa. 1983. Ge- 
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crabs from the Hampshire Basin. — Palaeontol- 
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135-150. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(l):53-69. 1996 

Systematics and distribution of the genus Calocarides 
(Crustacea: Decapoda: Axiidae) 

Brian Kensley 

Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — The nine currently recognized species of Calocarides are re- 
viewed, and two new species are described: C. capensis from the south-east 
Atlantic Ocean off the Cape of Good Hope, and C. macphersoni from the 
Atlantic Ocean off Namibia. Species of the genus are particularly characterized 
by the possession of dentate median, submedian, and lateral carinae of the 
carapace, weakly pigmented or unpigmented eyes, and robust, quite heavily 
sclerotized, tuberculate and/or spinose first pereopods. 



The approximately 100 described species 
of Axiidae show their greatest diversity in 
shallow tropical and subtropical seas, with 
several genera confined to the tropics, e.g., 
Coralaxius Kensley & Gore, 1982, Parax- 
iopsis De Man, 1905, Paraxius Bate, 1888, 
Scytoleptus Gerstaecker, 1856. A few gen- 
era, however, have colonized colder and 
deeper waters and show considerable diver- 
sity and broad geographic range, e.g., Cal- 
ocarides Wollebaek, 1908, and Eiconaxius 
Bate, 1888, the latter an inquiline of hex- 
actinellid sponges. Calocarides is geo- 
graphically widespread in continental shelf- 
slope regions, and contains cold-water in- 
habiting, soft-bottom burrowing species, 
some of which may be associated with 
zones of upwelling. The genus is reviewed 
here, two new species are described, and a 
number of taxonomic and distributional 
problems noted, relating to species previ- 
ously included in the genus. 

Abbreviations: cl — carapace length; 
HUMZ — Laboratory of Marine Zoology, 
Hokkaido University; ICMB — Instituto de 
Ciencias del Mar, Barcelona; MNHN — Mu- 
seum National d'Histoire Naturelle, Paris; 
SAM — South African Museum, Cape 
Town; UBZM — Zoological Museum, Uni- 
versity of Bergen; USNM — National Mu- 



seum of Natural History, Smithsonian 
Institution. 

Systematic Account 

Family Axiidae 

Genus Calocarides Wollebaek, 1908 

Calocaris (Calocarides) Wollebaek, 1908: 

3, 23. 
Axiopsis (Calocarides). — De Man, 1925:1, 

2, 6, 67, 71.— Balss, 1957:1579. 
Calocarides. — Bouvier, 1940:97. — Sakai & 

de Saint Laurent, 1989:4, 78.— Poore, 

1994:98. 
Euconaxius [laps. cal. for Eiconaxius Bate, 

1888] Trybom, 1904:384. 

Type species. — By subsequent designa- 
tion by Sakai & de Saint Laurent, 1989, Eu- 
conaxius coronatus Trybom, 1904. Type lo- 
cality: Skagerrak, North Sea, 230-500 m. 
Gender: Masculine. 

Diagnosis. — Gonochoristic. Carapace 
having small supraocular spine present, as 
part of lateral rostral series; post-cervical 
carina low, rounded, barely discernible in 
some species, lacking spines; rostrum at 
same level as anterior carapace, margins 
armed; median carina dentate; submedian 
carina dentate; lateral carina dentate. 

Eye reaching to about midlength of ros- 
trum, cornea weakly pigmented or unpig- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1 . — Branchial formula for Calocarides coron- 
atus. r = reduced. 



Maxillipeds 



Pereopods 



Exopod 1 1 1 — — — — — 

Epipod 111 1111 — 

Podobranch — — r rrr — — 

Arthrobranch — — 2 2222 — 

Pleurobranch — — — — — — — — 

Setobranch — — 1 11111 



merited; eyestalk longer than eye. Antennal 
scaphocerite elongate. Maxilla 2, scaphog- 
nathite having spinulose flagellum. Maxil- 
lipeds 1-3 with exopods and epipods. Pe- 
reopod 1, chelipeds asymmetrical, some- 
what sexually dimorphic. Propodi and dac- 
tyli of pereopods 3-5 slender, not 
expanded, dactyli simple. Pleopodal rami 
slender elongate; appendix interna present 
on pleopods 2-5. Pleopod 1 S absent; pleo- 
pod 1 $ uniramous, of 2 articles. Pleopod 
2 S with appendix masculina of single 
lobe. Uropod with lateral ramus with su- 
ture. Telson longer than wide, or at least as 
long as wide, with dorsal spines. 
Branchial formula: see Table 1. 
Species: 
Calocarides capensis, new species. Off Ta- 
ble Bay, South Africa, 1280 m. 
Calocarides coronatus (Trybom, 1904). 
Swedish and Norwegian fjords. North 
Sea, 80-700 m. 
Calocarides lev (Zarenkov, 1989). Gulf of 

California, 2000-2036 m. 
Calocarides longispinis (McArdle, 1901). 

Bay of Bengal, 549 m. 
Calocarides macphersoni, new species. Off 

Namibia, 300-480 m. 
Calocarides quinqueseriatus (Rathbun, 
1902). Off mouth of Columbia River, Or- 
egon, to southern California, 292-2013 m. 
Calocarides rostriserratus (Andrade & 
Baez, 1977). Off Chile, 320^00 m; off 
Peru, Gulf of Panama, 576-825 m. 
Calocarides rudolfi (Zarenkov, 1989). Off 
Angola, 490-510 m. 



Calocarides soyoi (Yokoya, 1933). Off 
northern Japan, 138-270 m. 

Calocarides spinulicauda (Rathbun, 1902). 
Off Bodega Head, California, 113 m; Pu- 
get Sound, 70-256 m; British Columbia, 
91-128 m. 

Calocarides vigila Sakai, 1992. New Zea- 
land, 74-146 m. 

Taxonomic remarks. — The species Cal- 
ocaris lev Zarenkov, 1989, described from 
a hydrothermal vent at 2000-2036 m in the 
Gulf of California, is almost certainly a spe- 
cies of Calocarides. The proportions and 
spination of the first chelipeds bear more 
resemblance to C. rostriserratus from 
Chile, Peru, and Panama, than to C. quin- 
queseriatus from southern California. Giv- 
en that this species was taken from greater 
depth than either of the two abovemen- 
tioned species, its relative isolation of the 
Gulf of California, and the lack of males, it 
seems prudent to await further material be- 
fore deciding on the status of this species. 

Calocaris rudolfi Zarenkov, 1989, from 
490-510 m off Angola is likewise a species 
of Calocarides, and bears some resem- 
blance to C. macphersoni (see Remarks un- 
der C. macphersoni). 

Although Sakai & de Saint Laurent 
(1989) tentatively ^\2iCe.d Axiopsis tenuicor- 
nis De Man, 1905, Axius habereri Balss, 
1913, Axius armatus Smith, 1881, and Ax- 
iopsis werribee Poore & Griffin, 1979, 
within Calocarides, none of these possess 
a majority of the features that define the 
genus. 

The specific identity of Calastacus quin- 
queseriatus and Axius spinulicauda amu- 
rensis, both recorded from Japan by Kob- 
jakova (1937) will remain uncertain until 
the material can be located and examined. 

Ecological and biological notes. — Little 
information exists for species of Calocari- 
des, except for C coronatus and C quin- 
queseriatus. Both of these species are 
known to burrow into soft bottom sedi- 
ments. In the case of C coronatus, these 
sediments are usually mud with a high 



VOLUME 109, NUMBER 1 



55 



(60%) silt content and a temperature of 6°- 
7.5°C. Calocarides coronatus overlaps in 
bottom distribution with the calocarid 
shrimp Calocaris macandreae, but domi- 
nates in deeper water (Brattegard 1966). 
For all the records of C. quinqueseriatus, 
the bottom sediments were recorded as ei- 
ther yellow or green mud, with tempera- 
tures ranging from 13°-16°C. 

Even less is known of the prey organisms 
of species of Calocarides. Calocarides 
macphersoni from Namibia was taken from 
the gut-contents of two bottom feeders, viz. 
the scorpaenid fish Helicolenus sp., and the 
ray Raja confundens, again suggesting that 
the species burrows in soft sediments. 

Regarding reproduction and fecundity, C. 
coronatus produces a about 65 eggs/female, 
with an egg diameter 15% of carapace 
length {n = 2). Calocarides quinqueseriatus 
produces about 180 eggs/female, with an 
egg diameter of about 7% of carapace length 
(n = 4). These egg sizes are in the high 
range for free-living axiids, suggesting that 
a fairly advanced larva hatches, having a rel- 
atively short planktonic larval lifespan. Spe- 
cies which produce thousands of eggs per 
female generally have much smaller eggs, 
e.g., Oxyrhynchaxius caespitosa with 16,000 
eggs/female, egg diameter 1.3% of carapace 
length; Axiopsis serratifrons with 4000 eggs/ 
female, egg diameter 2.7% of carapace 
length. The commensal species of Eiconax- 
ius by contrast, produce far fewer large eggs, 
8-30 eggs/female, with an egg diameter 16- 
38% of carapace length. Elofsson (1959) re- 
corded a 10 mm stage II larva (Gumey 
1942) of C coronatus from 600 m. 

Distribution. — From the depth ranges of 
the 1 1 species of Calocarides (see list of 
species), it is clear that they are continental 
shelf/slope dwellers, probably with a pref- 
erence for soft-sediment bottoms into which 
they can burrow. In several cases, the shelf/ 
slope habitat is also characterized by up- 
welling of deeper water, e.g., off the Cape 
of Good Hope and Namibia, off Pacific 
North America, off Chile, and in localized 
areas off Norway. 



The widespread geographical distribution 
of the species of Calocarides, in the Atlan- 
tic, Indian, and Pacific oceans (Fig. 1), 
along with the most northerly (C corona- 
tus) and most southerly (C vigila) records 
for the Axiidae, suggests a considerable age 
for the genus. If, as has been suggested, the 
axiid-like shrimps evolved from Tethyan 
ancestors (Kensley 1994), shallow- water 
precursors of Calocarides must have spread 
with tectonic plate movements and the 
opening of the present-day oceanic basins, 
at the same time penetrating to varying 
depths the waters of the continental shelves. 
The direction of this invasion from shallow 
to deeper shelf waters may be reflected in 
that some eye-pigment is retained in some 
species (e.g., C soyoi) while other species 
completely lack eye-pigment (e.g., C. ca- 
pensis). 

Calocarides capensis, new species 
Fig. 2 

Calastacus longispinis. — Stebbing, 1910: 

367, non McArdle, 1901. 
Calocaris {Calastacus) longispinis. — De 

Man, 1925:8 (part), non McArdle, 

1901.— Barnard, 1950:503, fig. 93 d-f. 
Calocaris longispinis. — Kensley, 1981:30, 

non McArdle, 1901. 
Calocarides coronatus. — Sakai & de Saint 

Laurent, 1989:79, non Trybom, 1904. 

Material examined. — Holotype, SAM- 
A940, ovigerous 9 cl 19.0 mm (carapace 
damaged), R/V Pieter Faure 40 miles 
N79°E of Table Mountain, Cape of Good 
Hope, South Africa, 1280 m. 

Diagnosis. — Carapace surface minutely 
pitted, especially posteriorly; rostrum with 
5-6 lateral teeth; median carina well 
marked, unarmed; submedian carina bear- 
ing 1 or 2 spines; lateral carina with 3 or 4 
spines. Abdominal pleura ventrally round- 
ed, pleuron 2 widest. Telson with 1 pair of 
small spines dorsally, 3^ small mobile 
spines at posterolateral corner, posterior 
margin gently convex between posterolat- 
eral and median spines. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1 . Distribution of species of Calocarides. 



Maxilliped 3, merus with 4 spines on 
posterior margin; carpus with single small 
distal spine. Pereopod 1, larger cheliped, is- 
chium with single strong distal spine; merus 
with strong spine at midlength of posterior 
margin, few small tubercles in proximal 
half, anterior margin with 5 distal spines; 
carpus with few low tubercles on anterior 
margin and few small tubercles on lateral 
surface; propodal palm slightly longer than 
fingers, outer surface with tiny scattered 
acute tubercles and strong ridge near pos- 
terior margin reaching almost to tip of fixed 
finger, anterior margin with about 1 1 strong 
spines, cutting margin of finger finely 
toothed, with large proximal tubercle; dac- 
tylus with few tubercles on anterior margin, 
cutting edge finely toothed, with proximal 
tubercle. Smaller cheliped, ischium with 
strong distal spine; merus with strong spine 
at midlength of posterior margin, anterior 
margin with 3 distal spines; carpus with few 
tubercles on anterior margin and lateral sur- 
face; propodal palm longer than fingers, lat- 



eral surface with few scattered small tuber- 
cles and ridge near posterior margin, ante- 
rior margin with about 1 1 spines; fixed fin- 
ger and dactylus as in larger cheliped. 
Pereopod 2, ischium with few small spines 
and strong distal spine on posterior margin; 
merus with 3 widely spaced spines on pos- 
terior margin. Pleopod 1 consisting of sin- 
gle slender elongate ramus. Pleopods 2-5 
each with appendix interna on mesial mar- 
gin of endopod. Lateral uropodal ramus 
with single small spine on outer margin; su- 
ture bearing 4-8 irregular spines; mesial ra- 
mus with 4 spines on middorsal ridge. 

Remarks. — See 'Remarks' after Calocar- 
ides macphersoni. 

Etymology. — The specific name derives 
from the type locality, the Cape of Good 
Hope. 

Calocarides coronatus (Trybom, 1904) 

Euconaxius coronatus Trybom, 1904:384— 
390, pi. 20, figs. 1-10, 13, 14, pi. 21, figs. 
1-8. 



VOLUME 109, NUMBER 1 



57 




Fig. 2. Calocarides capensis, new species: A, Carapace in lateral view, scale = 5 mm; B, Anterior carapace 
in dorsal view; C, Abdomen in lateral view; D, Telson in dorsal view; E, Pleopod 1, female; F, Pleopod 2, 
female; G, Pereopod 1, smaller cheliped; H, Pereopod 1, larger cheliped; I, Maxilliped 3; J, Pereopod 2. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Calocaris (Calocarides) coronatus. — Wol- 
lebaek, 1908:3, 5, 13, 23. 

Axiopsis {Calocarides) coronatus. — De 
Man, 1925:6, 67, 71. 

Calocarides coronatus. — Bjorck, 1913:7. — 
Balss, 1925:209.— Poulsen, 1940:208, 
216, fig. 4.— Bouvier, 1940:97.— Chris- 
tiansen, 1955:1^; 1972:40, fig. 45.— 
Elofsson, 1959:3-9, figs. 1-20.— Bratte- 
gard, 1966:45-52.— Sakai «fe de Saint 
Laurent, 1989:4, 80, 101, fig. 20. 

Euconaxius crassipes Trybom, 1904:390- 
391, pi. 20, figs. 11, 12. 

Calocaris {Calocarides) crassipes. — Wol- 
lebaek, 1908:3, 5-13, 17, 23, pi. 1-7. 

Axiopsis {Calocarides) crassipes. — De 
Man, 1925:6, 67, 71. 

Material examined. — UBZM 50400, S 
cl 14.8 mm, Sorfjord, Norway, 60°11'N, 
6°34'E, 589-600 m.— UBZM 49598, 4 S 
cl 13.5, 13.9, 15.5, 15.5 mm, Kvinnherad- 
fjord, Norway, 59°56'N, 5°45'E, 512 m.— 
UBZM 49595, 2 cl 14.0 mm, Kvinnherad- 
fjord, Norway, 59°59'N, 5°53'E, 660-670 
m.— UBZM 50461, ovigerous 9 cl 14.4 
mm, Osafjord, Norway, 60°40'N, 6°55'E, 
300.— UBZM 49589, Alfjord, Norway, 
59°4rN, 5°34'E, 450-465 m.— UBZM, 
ovigerous $ cl 16.1 mm, Fusafjord, Nor- 
way, 60°12'N, 5°45'E, 240 m.— UBZM 
49591, S cl 15.1, 2 cl 15.2 mm, Bomla- 
fjord, Norway, 59°40'N, 5°22'E, 360 m. 

Distribution. — From Malangenfjord, 
northern Norway, to southwestern Sweden, 
80-700 m. 

Diagnosis. — Carapace surface smooth. 
Rostrum with 4 pairs of lateral teeth, con- 
tinuous with lateral carina bearing 3 teeth; 
median carina with 1 tooth anterior to tu- 
bercle; submedian carina bearing 2 teeth. 
Abdominal pleuron 1 ventrally narrowly 
rounded; pleuron 2 broadly rounded; pleura 
3-5 rounded, with 1 or 2 small teeth on 
anterior margin. Telson with 5 or 6 lateral 
teeth, 2 mobile spines at posterolateral an- 
gle; posterior margin evenly convex, with 
median tooth; 1 or 2 small spines proxi- 
modorsally on rounded ridge. 



Antennal scaphocerite reaching distal 
margin of peduncle article 4. Maxilliped 3, 
merus with 4 spines on posterior margin, 3 
more distal spines large; carpus with small 
distal spine on posterior margin. Pereopod 
1, larger cheliped: ischium with row of 
acute tubercles on posterior margin, distal- 
most a large spine; merus with posterior 
surface expanding distally, mesial and lat- 
eral margin defined by row of small acute 
tubercles, mesial margin with strong spine 
at about midlength, anterior margin bearing 
1 strong spine and few blunt tubercles; car- 
pus bearing small tubercles dorsodistally; 
fingers slightly more than half length of 
propodal palm, latter with lateral and most 
of mesial surface tuberculate, fixed finger 
with triangular tooth at about midlength of 
cutting edge; fingers widely gaping, dacty- 
lus downcurved, strongly tuberculate, with 
1 or 2 stronger tubercles on cutting edge. 
Smaller cheliped: ischium, merus, and car- 
pus as in larger cheliped; propodal palm 
with lateral and mesial surface tuberculate, 
tubercles along anterior margin forming 
spines, fingers about % length of palm, 
fixed finger with blunt proximal triangular 
tooth followed by regularly spaced small 
teeth; dactylus with row of spines along an- 
terior margin, cutting edge with regularly 
spaced small teeth. Pereopod 2, ischium 
and basis each with single posterodistal 
spine; merus with 3 evenly spaced spines 
on posterior margin. Lateral uropodal ra- 
mus with about 18 spines along suture, 2 
small distal spines on lateral margin; mesial 
ramus with single distolateral spine, dorsal 
ridge bearing 5 or 6 spines. 

Calocarides macphersoni, new species 
Fig. 3 

Calastacus longispinis. — Macpherson, 
1983:45, fig. 26; 1991:405, non Mc- 
Ardle, 1901. 

Material examined. — Holotype, USNM 
243561, S cl 20.4 mm, RA^ Benguela II sta 
66, off Namibia, 20°55'S, 12°23'E, 475- 
480 m, 9 Sep 1980.— Paratypes, USNM 



VOLUME 109, NUMBER 1 



59 




Fig. 3. Calocarides macphersoni, new species: A, Lateral view, scale = 5 mm; B, Telson and right uropod 
in dorsal view; C, Pereopod 2; D, Maxilliped 3; E, Pereopod 1, larger cheliped; F, Pereopod 1, smaller cheliped. 



243562, S cl 17.8 mm, RA^ Benguela II sta 
66, off Namibia, 20°55'S, 12°23'E, 475- 
480 m, 9 Sep 1 980.— Paratype, ICMB, R/ 
V Benguela II sta P-37, S cl 21.5 mm, off 
Namibia, 21°33'S, 12°39'E, 380-390 m, 
from stomach of scorpaenid fish Helicolen- 



us sp. — Paratypes, ICMB, KTV Benguela II 
sta P-66, 2 S c\ 20.3 mm, one damaged, 
off Namibia, 20°55'S, 12°23'E, 475^80 
m. — Paratypes, ICMB, RA^ Benguela IV 
sta P-95, 2 S cl 21.0 mm, 21.2 mm, off 
Namibia, 20°42'S, 12°14'E, 410^17 m. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



one specimen from stomach of ray Raja 
confundens, one from stomach of scorpaen- 
id fish Helicolenus sp. 

Diagnosis. — Rostrum with 2-3 pairs of 
lateral teeth; median carina with single tooth 
anterior to tubercle, entire posterior to tu- 
bercle; submedian carina poorly defined, 
with 1 or 2 teeth; lateral carina with 3-5 
teeth posterior to supraocular. Carapace sur- 
face smooth. First abdominal pleuron ven- 
trally narrowly rounded, pleura 2-6 broadly 
rounded, small spine on anterior margin of 
pleura 3-6. Telson with 5 small fixed spines 
in posterior half of lateral margin, ?>—^ small 
mobile spines mesial to posteriormost lateral 
spine; posterior margin gently convex be- 
tween lateral spine and median spine; dorsal 
surface with one pair of small fixed spines. 

Maxilliped 3, posterior margin of ischi- 
um unarmed; merus with 3-4 strong distal 
spines and 2-3 tiny more proximal spines 
on posterior margin; carpus with small pos- 
terodistal spine. Pereopod 1: larger chela, 
ischium with row of small denticles and 
single strong spine along posteromesial 
margin, small distal spine on anterior mar- 
gin; merus with posteromesial surface tu- 
berculate, having single strong tooth at 
about midlength, anterior margin with 
about 5 small spines and several small tu- 
bercles; outer and upper surface of carpus 
evenly tuberculate; propodal palm about Vg 
longer than fingers, entire surface evenly 
and fairly densely tuberculate, posterior 
margin with tuberculate carina extending 
onto fixed finger, anterior margin with row 
of somewhat larger tubercles; fingers prox- 
imally widely gaping, distally meeting, 
fixed finger with proximal half of cutting 
edge having few low rounded tubercles, 
distal half straight, bearing numerous small 
rounded teeth; surface of dactylus tubercu- 
late, cutting edge as in fixed propodal fin- 
ger. Smaller chela, ischium with few small 
spines and single strong spine on posterior 
margin, single distal spine on anterior mar- 
gin; merus with row of small tubercles 
along posteromesial margin with 1 or 2 
strong spines at about midlength, row of 



small tubercles along posterolateral margin, 
anterior margin with row of irregularly 
large and small spines and tubercles; outer 
and upper surface of carpus fairly densely 
tuberculate; propodal palm about '/g longer 
than fingers, posterolateral carina feebly tu- 
berculate, extending almost to apex of fixed 
finger, outer and upper surfaces tuberculate, 
anterior margin with double row of large 
and small spines and tubercles; cutting mar- 
gins of fixed propodal finger and dactylus 
only slightly excavate proximally, remain- 
der of margin bearing row of small evenly- 
spaced teeth. Pereopod 2, posterior margin 
with row of small spines and single strong 
distal tooth; posterior margin of merus with 
2-4 spines. Lateral uropodal ramus with 2 
or 3 small distal spines on lateral margin, 
single mobile spine mesial to distalmost, 
transverse suture armed with row of 13-15 
small spines; mesial ramus with single dis- 
tal spine, dorsal ridge with irregular row of 
about 8 small spines. 

Remarks. — From McArdle's (1901) de- 
scription, and Alcock & McArdle's illustra- 
tions (1902, pi. 57, figs. 2, 2a), and De 
Man's (1925:118) key, the Bay of Bengal 
species Calastacus longispinis lacks spines 
on the anterior median carina, has five 
spines on the free lateral margins of the ros- 
trum, and three spines on the submedian ca- 
rina. The antennal scaphocerite reaches al- 
most to the end of peduncle article 4. The 
single female lacked both chelipeds of pe- 
reopod 1 . Given that the Namibian material 
has a strong spine anterior to the tubercle 
on the median carina, and that the antennal 
scaphocerite reaches only to the distal third 
of peduncle article 4, it seems unlikely that 
the Atlantic and Indian Ocean specimens 
are the same species, especially considering 
the considerable geographical separation. 

The species described above as Calocar- 
ides capensis, differs from C. macphersoni 
in having more slender and less granular 
chelipeds of pereopod 1, with relatively 
more elongate spines on the anterior margin 
of the propodal palm; in having fewer but 
stronger distal spines on the anterior margin 



VOLUME 109, NUMBER 1 



61 



of the merus of pereopod 1; in having far 
fewer spines on the suture of the lateral ra- 
mus of the uropod; and in having a more 
elongate telson. 

Major differences between C. coronatus- 
and C. macphersoni are seen in the size (cl 
ovigerous 9 13.5-16.1 mm vs. cl ovigerous 
2 17.8-25.0 mm resp.), and in the more 
spinose upper margin of the merus in C. 
macphersoni. 

The species described as Axius (Neaxius) 
laevis by Bouvier (1915) from 698 m off 
Cape Bojador, Spanish Sahara, West Africa, 
(holotype, MNHN, ? cl 5.1 mm) was syn- 
onymized with C. coronatus by Sakai & de 
Saint Laurent (1989:81, fig. 21). The latter 
authors illustrated the single damaged spec- 
imen of C. laevis, the caption of their illus- 
tration erroneously referring to Eiconaxius 
laevis. Given the damaged condition of the 
specimen, and the fact that it lacks spines 
on the submedian carina, has an antennal 
scaphocerite shorter than in C. coronatus, 
has pereopod 1 lacking the fine tubercula- 
tion of the propodus and the anterior spi- 
nation of the merus of C. coronatus, and a 
telson longer and narrower than in C. co- 
ronatus, it seems unlikely that the West Af- 
rican species is synonymous with C. coron- 
atus, or indeed, that it is a species of Cal- 
ocarides. From the figure of Calocarides 
rudolfi (Zarenkov, 1989, fig. 3) from An- 
gola, several differences in spination sug- 
gests that it differs from C. macphersoni. In 
C. rudolfi, the merus of maxilliped 3 bears 
more elongate and numerous spines, and 
the merus of both chelae of pereopod 1 has 
larger and more numerous anterodistal and 
posterodistal spines than in the Namibian 
species; the carpus of pereopod 2 bears a 
posterodistal spine, and the telson bears two 
pairs of fixed dorsal spines, while C. mac- 
phersoni lacks a carpal spine on pereopod 
2 and the telson bears a single pair of fixed 
spines. Posterolateral mobile spines on the 
telson were not illustrated for the Angolan 
species; C. macphersoni bears four such 
spines on each side. 

Etymology. — The species is named for 



carcinologist Dr. Enrique Macpherson of 
Barcelona, Spain, who made the material 
from Namibia available for study. 

Calocarides quinqueseriatus 

(Rathbun, 1902) 

Figs. 4, 5 

Calastacus quinqueseriatus Rathbun, 1902: 

887.— Schmitt, 1921:113, fig. 76.— 

Balss, 1925:209. 
Calocaris {Calastacus) quinqueseriata. — 

De Man, 1925:8, 118. 
Calocarides quinqueseriatus. — Sakai & de 

Saint Laurent, 1989:79, 103. 

Material examined. — Syntypes, USNM 
25240, 9 c? cl 15.5-22.7, 2 ovigerous 9 cl 
15.4, 18.2, 3 2 cl 17.5-19.4, 19.4, RJV Al- 
batross sta 3196, San Luis Obispo Bay, 
California, green mud, 366 m. — USNM 
28321, 5 c?, cl 13.5-21.6, 1 ovigerous 9 cl 
16.9 mm, 9 9 cl 15.1-20.5, R/Y Albatross 
sta 2909, Santa Barbara Channel, Califor- 
nia, 375 m, green mud. — USNM 152525, 
22 S cl 18.9-26.0, 9 ovigerous 9 20.0- 
23.3, 11 9 cl 21.0-23.0, R/Y Albatross sta 
4436, San Miguel Island, California, 483- 
496 m, green mud.— USNM 28325, 16 S , 
cl 11.3-18.0, 1 ovigerous 9 cl 24.0, 7 9 cl 
10.2-19.2, RA^ Albatross sta 3195, San 
Luis Obispo Bay, California, 461 m, green 
mud.— USNM 28236, 5 d cl 14.8-24.0, 4 
9 cl 13.0-22.5, RA^ Albatross sta 3198, off 
Point Conception, California, 508 m, green 
mud.— USNM 28327, 7 c? cl 12.9-20.5, 3 
ovigerous 9 cl 19.0-20.0,6 9 cl 10.3-19.5, 
RA^ Albatross sta 3199, Santa Barbara 
Channel, California, 426 m, green mud. — 
USNM 28232, 6 cl 18.0, 2 9 cl 21.0, 21.9 
R/V Albatross sta 3187, off Point Sur, Cal- 
ifornia, 545 m, brown and grey ooze. — 
USNM 28320, S cl 20.3, 9 cl 17.2, R/V 
Albatross sta 2892, off Point Conception, 
California, 520 m, yellow mud. — USNM 
28528, 2 S, cl 18.1, 24.1, RA^ Albatross 
sta 3200, Santa Barbara Channel, Califor- 
nia, 485 m, green mud.— USNM 28319, 1 
S cl 19.0, 1 9 cl 12.1, RA^ Albatross sta 
2891, off Point Conception, Cahfornia, 426 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 4. Calocarides quinqueseriatus: A, Lateral view, scale = 10 mm; B, Pereopod 1, larger cheliped; C, 
Pereopod 1, smaller cheliped; D, Telson and right uropod in dorsal view; E, Pleopod 2, male; F, Pleopod 3, 
male; G, Pleopod 1, female; H, Pleopod 2, female. 



m, brown ooze.— USNM 120119, 9 cl 
18.2, RA*" Commando sta, mouth of Colum- 
bia River, Oregon, 732 m.— USNM 28324, 
S cl 17.5, 9 cl 15.0, RA^ Albatross sta 
3193, off San Simeon Bay, California, 293 
m, green mud. — USNM 28329, ovigerous 
9 cl 19.5, RA^ Albatross sta 3201, Santa 
Barbara Channel, California, 512 m. — 
USNM 28322, S cl 17.4, RA^ Albatross sta 
2979, off Anacapa Island, California, 710 



m, green mud.— USNM 53021, 9 cl 16.2, 
RA^ Albatross sta 4425, off San Nicolas Is- 
land, California, 2013 m, green mud, fine 
sand, globigerina ooze. — USNM 155735, 
S cl 20.5, RA^ Albatross sta 4387, off San 
Diego, California, 1830 m, green mud. — 
USNM 243350, ovigerous 9 damaged, R/ 
V Commando sta, SW mouth of Columbia 
River, Oregon, 1556 m.— USNM 243351, 
S cl 18.4, RA^ Commando sta, SW mouth 



VOLUME 109, NUMBER 1 



63 












Fig. 5. Calocarides quinqueseriatus, variation in rostral and anterior carapace spination: A, Syntype, ovi- 
gerous female, San Luis Obispo Bay, California, cl 15.4 mm; B, Syntype, ovigerous female, San Luis Obispo 
Bay, California, cl 18.2 mm; C, Ovigerous female, San Miquel Is., California, cl 22.0 mm; D, Ovigerous female, 
San Miguel Is., California, cl 23.3; E, Female, Baja California, cl 12.0 mm; F, Female, Baja California, 20.7 
mm; G, Male, Columbia River mouth, Oregon, 18.2 mm; H, Syntype, male, San Luis Obispo Bay, California, 
cl 19.1 mm; I, Syntype, male, San Luis Obispo Bay, California, cl 20.9 mm; J, Male, San Miguel Is., California, 
24.2 mm; K, Male, San Miguel Is., California, cl 26.0 mm; L, Male, Columbia River, Oregon, cl 27.2 mm. 



of Columbia River, Oregon, 1373 m. — 
USNM 243352, 9 damaged, RA^ Comman- 
do sta, SW mouth of Columbia River, Or- 
egon, 1556 m.— USNM 243353, S cl 17.3, 
2 9 damaged, RA'^ Commando sta, SW 
mouth of Columbia River, Oregon, 1922 
m.— USNM 243354, 9 damaged, R/V 
Commando sta, SW mouth of Columbia 
River, Oregon, 1556 m.— USNM 155737, 
9 cl 20.0, RA^ Albatross sta 5675, San 
Cristobal Bay, Baja California, 520 m. — 
USNM 155736, 2 c? cl 19.8-21.0, RA^ Al- 
batross sta 4433, off Santa Rosa Island, 



California, 445 m, green mud. — USNM 
243355, 2 (? cl 16.5, 2 9 cl 13.8-15.0, R/ 
V Albatross sta 4351, NE of Point Loma, 
California, 774 m, soft green mud. 

Diagnosis. — Carapace smooth; 5-6 pairs 
rostral teeth including small supraocular; 
median carina with 1-4 spines anterior to 
tubercle, 1-2 posterior; 1-5 spines on sub- 
median carina; 3-7 spines on lateral carina. 
Abdominal pleuron 1 with ventral spine 
usually present; pleuron 2 ventrally round- 
ed; pleura 3-6 rounded with spine on an- 
terior margin. Telson with lateral margin 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



having 5-6 small spines; 1 pair spines on 
dorsal surface. 

Maxilliped 3, ischium with about 10 
small spines on posterior margin; merus 
with 5-6 small proximal spines, 3 large dis- 
tal spines on posterior margin. Pereopod 1, 
larger (broader) cheliped: ischium with 1 
strong distal and several small proximal 
spines on posterior margin; merus with in- 
ner and outer row of tiny tubercles flanking 
posterior row of about 17 strong spines, 
outer surface distally with few scattered 
tiny tubercles, upper (anterior) margin with 
irregular row of spines, distal spines largest; 
carpus with outer surface and distal margin 
bearing scattered acute tubercles; upper 
propodal palm subequal in length to fingers, 
posterior margin of palm and fixed finger a 
denticulate carina, upper (anterior) margin 
with about 14 strong spines, mesial and lat- 
eral surfaces of palm fairly densely granu- 
lar/tuberculate, fixed finger with broad tu- 
bercle on proximal cutting edge, distally 
denticulate; dactylus cutting edge with 
proximal tubercle, distally denticulate, up- 
per margin with several proximal spines. 
Pereopod 1, smaller (narrower) cheliped: is- 
chium, merus, and carpus as in larger che- 
liped; upper propodal palm subequal in 
length to fingers, tuberculation as in larger 
cheliped, cutting edge of fixed finger with 
few small tubercles proximally, remainder 
of edge finely denticulate; dactylus, cutting 
edge finely denticulate, with about 2 spines 
on upper margin. Pereopod 2, ischium with 
6 small spines on posterior margin; merus 
with 5 small proximal spines, single strong 
distal spine. Pleopod 1 in female a slender 
elongate ramus; pleopod 2 in male with 
rod-shaped appendix masculina bearing dis- 
tal setae, slightly shorter than appendix in- 
terna; pleopods 2-5 in female, 3-5 in male 
with appendix interna at about proximal 
third of mesial margin of endopod. Uropod 
with lateral ramus having 2 small distal 
spines on lateral margin, single mobile sub- 
marginal spine; about 14 spines along su- 
ture; mesial ramus with single distolateral 
spine, 4 spines on dorsal surface. 



Size ranges. — Male, maximum carapace 
length 26.0 mm; ovigerous females, cara- 
pace length 15.4-24.0 mm. 

Remarks. — There is considerable varia- 
tion in the spination of the anterior carapace 
(Fig. 5) and the chelipeds of pereopod 1 of 
this species, often made more marked either 
by injuries to the chelae that have healed, 
or by loss and regeneration of the chelipeds, 
resulting in a range of cheliped proportions 
and armature. 

The majority of specimens have been 
taken from soft bottom sediments, especial- 
ly from green mud. 

The species recorded as Calastacus quin- 
queseriatus from 1150 m off Japan (Kob- 
jakova 1937:142, pi. 2, fig. 8) appears to 
have the first pereopod chelae relatively 
shorter, more robust, and less granular than 
in true C. quinqueseriatus. It thus seems un- 
likely that the Japanese species is conspe- 
cific with the California/Oregon species. 

Calocarides rostriserratus 

Andrade & Baez, 1977 

Fig. 6 

Calastacus rostriserratus Andrade & Baez, 

1977:65, fig. 1. 
Calocarides quinqueseriatus. — Sakai & de 

Saint Laurent, 1989:78, 79, 103, non 

Rathbun, 1902. 

Material examined. — USNM 173366, 4 
S cl 15.3 mm, 15.3 mm, 16.3 mm, 17.0 
mm, 2 9 cl 14.5 mm, 15.9 mm, off Tum- 
bes, Peru, 3°33.8'S, 81°01.4'E, 576 m, 28 
Aug 1979.— USNM 243392, $ cl 16.1 mm, 
RA^ Gilliss sta GS-22, Gulf of Panama, 
7°28'N, 79°12'W, 825 m, 18 Jan 1972. 

Diagnosis. — Carapace smooth; rostrum 
with 5-6 pairs of lateral teeth including 
small supraocular; 2 median carina teeth 
plus tubercle; 4-5 teeth on submedian ca- 
rina; 4-6 teeth on lateral carina. Abdominal 
pleura 2-5 ventrally rounded. Telson with 
2-3 pairs small lateral spines, 2 small distal 
mobile spines at posterolateral angle, 2 
pairs spines on dorsal surface. 

Maxilliped 3, ischium with 5 small prox- 



VOLUME 109, NUMBER 1 



65 




Fig. 6. Calocarides rostriserratus: A, Lateral view, scale = 5 mm; B, Anterior carapace in dorsal view; C, 
Telson and right uropod in dorsal view; D, Pereopod 1, larger cheliped; E, Pereopod 1, smaller cheliped; F, 
Pleopod 1, female; G, Pleopod 2, female; H, Pleopod 2, male; \, Pleopod 3, male. 



imal teeth on posterior margin, distal mar- 
gin entire; merus with 2 strong distal teeth 
on posterior margin. Pereopod 1, larger 
(broader) cheliped, ischium with about 5 
small proximal and single strong distal 
spine on posterior margin; merus with 5 
distal spines on upper (anterior) margin, 
posterior margin with single row of spines 
of varying length, those at about midlength 
longest; carpus with several (6-7) small 
spines on upper surface; upper propodal 



palm 1.2 times length of fingers, bearing 
row of 8 strong spines, lower border con- 
sisting of flattened distally narrowed ridge 
having dentate margins, medial and lateral 
surfaces bearing moderately dense scattered 
conical tubercles, cutting edge of fixed fin- 
ger having rounded proximal cusp; dactylus 
with upper margin bearing about 8 strong 
spines, cutting edge having rounded proxi- 
mal cusps, distally finely denticulate. Pereo- 
pod 1, smaller (narrower) cheliped, ischium 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



with 2-3 small proximal and single strong 
distal spine on posterior margin; merus with 
5-6 strong distal spines on anterior (upper) 
margin, single row of spines along posterior 
(lower) margin, those at about midlength 
longest; carpus with 8-10 spines on upper 
surface; upper propodal palm about 1.2 
times length of fingers, bearing irregular 
row of about 12 spines, lower border 
formed by flattened ridge with dentate mar- 
gins, inner and outer surfaces bearing scat- 
tered acute conical tubercles; dactylus with 
several small spines on upper surface, cut- 
ting edge more or less evenly denticulate. 
Pereopod 2, basis with single spine on pos- 
terior surface; ischium with proximal and 
one or 2 stronger distal spine on posterior 
margin; merus with 3 or 4 spines in mid- 
region of posterior margin. Pleopod 1 in fe- 
male a slender elongate ramus; pleopod 2 
in male with rod-shaped appendix mascu- 
lina bearing distal setae, noticeably shorter 
than appendix interna; pleopods 2-5 in fe- 
male, 3-5 in male with appendix interna at 
about proximal third of mesial margin of 
endopod. Uropod with lateral ramus having 
2-3 small lateral spines plus distal submar- 
ginal mobile spine; about 13 spines along 
suture; mesial ramus with single distal mar- 
ginal spine, 4-5 spines on upper surface. 

Remarks. — In spite of Sakai & de Saint 
Laurent's (1989:80) statement that C. ros- 
triserratus is probably a synonym of C. 
quinqueseriatus, Calocarides rostriserratus 
differs from C. quinqueseriatus, its geo- 
graphically nearest congener, in several eas- 
ily seen features: in having two pairs of 
dorsal telsonic spines (one pair in C. quin- 
queseriatus), and two or three lateral tel- 
sonic spines (five or six in C quinqueser- 
iatus); and in having the chelae of pereopod 
1 broader and less elongate, and with spines 
only in the distal half of the anterior margin 
of the merus (full anterior margin of merus 
bearing spines in C. quinqueseriatus). 

Calocarides soyoi (Yokoya, 1933) 

Axius soyoi Yokoya, 1933:49, fig. 25. — 
Horikoshi et al., 1982:30, 33, 38, 39, 120, 
145, 168. 



Axiopsis {Axiopsis) soyoi. — Sakai, 1987: 
303. 

Calocarides soyoi. — Sakai & de Saint Lau- 
rent, 1989:4, 83, 103.— Kensley & Ko- 
mai, 1992:81, fig. 1. 

Material examined. — USNM 243563, 2 

2 cl 12.0 mm, 15.0 mm, ovigerous 9 cl 
17.9 mm, off Fukushima Prefecture, Japan, 
37°04'N, 141°31.7'E, 270 m.— USNM 
243564, 6 cl 14.2 mm, ovigerous ? cl 18.8 
mm, 9 cl 11.8 mm, off Hachinohe, Aomori 
Prefecture, Japan.— HUMZ-C990, 3 $ cl 
11.3 mm, 14.1 mm, 13.0 mm, off Hachi- 
nohe, Aomori Prefecture, Japan, 42°52.6'N, 
145°22.2'E, 152 m.— HUMZ-C994, $ cl 
16.9 mm, off Fukushima Prefecture, Japan, 
37°17.3'N, 141°21.4'E, 141 m.— HUMZ- 
C998, 3 (? cl 12.4 mm, 15.0 mm, 15.3 mm, 
off Fukushima Prefecture, Japan, 37°04'N, 
14r31.7'E, 270 m. 

Remarks. — This species was recently re- 
described and figured by Kensley & Komai 
(1992). 

Calocarides spinulicauda 

(Rathbun, 1902) 

Fig. 7 

Axius spinulicauda Rathbun, 1902:886; 

1904:149, fig. 90. 
Axiopsis spinulicauda. — Schmitt, 1 92 1 : 1 1 1 , 

fig. 74.— Butler, 1961:60, figs. 1, 2, pi. 1. 
Axiopsis (Axiopsis) spinulicauda. — De 

Man, 1925:6, 67, 69. 
Acanthaxius spinulicaudus. — Sakai & de 

Saint Laurent, 1989:4, 66, 103. 

Material examined. — Holotype, USNM 
25239, ? cl 15.5 mm, RA^ Albatross sta 
3172, off Bodega Head, California, 113 m. 

Diagnosis. — Carapace surface smooth; 
rostrum with 5 (left) and 6 (right) lateral 
teeth, including supraocular; median carina 
barely reaching onto base of rostrum, with 

3 spines anterior of tubercle, entire poste- 
rior to tubercle; postcervical median carina 
obsolete except close to posterior margin of 
carapace; submedian carina short, entire; 
lateral carina entire posterior to supraocular 



VOLUME 109, NUMBER 1 



67 





Fig. 7. Calocarides spinuUcauda: A, Carapace in lateral view, scale = 5 mm; B, Abdomen in lateral view; 
C, Anterior carapace in dorsal view; D, Telson and right uropod in dorsal view. 



spine of rostral series. First abdominal pleu- 
ron ventrally narrowed apically rounded 
with tiny denticle; pleura 2-6 broadly 
rounded. Telson with lateral margin bearing 
larger fixed anterior spine, 4-5 small more 
posterior fixed spines; posterior margin 
evenly convex; submedian longitudinal 
ridge bearing 3 fixed spines, pair of fixed 
anterior spines mesial to longitudinal ridge 
present. 

Maxilliped 3, ischium with 1 or 2 tiny 
denticles on posterior margin; merus with 3 



distal spines on posterior margin. Pereopod 
1, both chelipeds missing. Pereopod 2, me- 
rus with 2 or 3 well separated spines on 
posterior margin. Lateral uropodal ramus 
with 7 small distal spines on lateral margin, 
transverse suture with row of 11-12 small 
spines; mesial ramus with 2 strong spines 
on lateral margin, dorsal ridge with 7 or 8 
small fixed spines. 

Remarks. — Rathbun (1902), in her de- 
scription of A. spinuUcauda, noted that the 
chelae of the first pereopod were missing. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Butler (1961) figured both chela of pereo- 
pod 1, from material from British Colum- 
bia. While the lateral and mesial faces of 
the propodal palm do not appear to be tu- 
berculate, as is characteristic of Calocari- 
des, the spination of the dactyli, propodi, 
carpi, and meri are characteristic of this ge- 
nus. Calocarides spinulicauda, however, 
disagrees with Calocarides in lacking spi- 
nation on the lateral and submedian carinae. 
Sakai & de Saint Laurent (1989) placed this 
species in their new genus Acanthaxius, 
which is characterized by a tendency to 
having a double cornea, and a lack of pleu- 
robranchs, neither of which are seen in C. 
spinulicauda. With only the female holo- 
type available for examination, the nature 
of the male pleopods and the first pereopods 
being uncertain, unambiguous generic 
placement is almost impossible. 

Axius spinulicauda amurensis Kobjako- 
va, 1937, (see also Derjugin & Kobjakova 
1935:142), described from the Sea of Japan, 
while clearly not a subspecies of the Cali- 
fornian Calocarides spinulicauda Rathbun, 
1902, is difficult to place generically, given 
the single small figure and the abbreviated 
description. 

Acknowledgments 

I thank Dr. Enrique Macpherson of the 
Instituto Ciencias del Mar, Barcelona, Mrs. 
Michelle van der Merwe of the South Af- 
rican Museum, and Dr. A. Fosshagen of the 
Zoological Museum, Bergen, for the loan 
of material. The paper was considerably im- 
proved by the comments and suggestions of 
Dr. Rafael Lemaitre and anonymous re- 
viewers, to whom I am very grateful. 

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 

109(1 ):70-74. 1996 

A new species of the axiid shrimp genus Acanthaxius from the 
Caribbean (Crustacea: Decapoda: Thalassinidea) 

Brian Kensley 

Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — Acanthaxius kirkmilleri is described from slope depths (420-440 
m) in the Caribbean Sea off Anguilla, and is the first unambiguous record of 
the genus in the Atlantic. The species is characterized by a slender rostrum 
longer than the eyestalks, highly spinose and relatively slender chelipeds of 
pereopod 1, and seven spines on the suture of the lateral uropodal ramus. 



A single female specimen of a distinctive 
axiid shrimp was collected by the RA'^ John 
Elliott Pillsbury in 1969, off Anguilla. The 
specimen is described here, as part of a se- 
ries of papers documenting the diversity of 
the Axiidae in the western Atlantic, and is 
deposited in the collections of the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM). 

Family Axiidae 

Genus Acanthaxius 

Sakai & de Saint Laurent, 1989 

Acanthaxius Sakai & de Saint Laurent, 
1989:4, 12, 14, 66.— Poore, 1994:98 
[key].— Sakai, 1994:192. 

Type species. — By original designation, 
Axiopsis (Axiopsis) pilocheira Sakai, 1987. 
Type locality: off Honshu Island, Japan, 
360 m. Gender: Masculine. 

Diagnosis. — Gonochoristic. Rostrum at 
same level as anterior carapace, narrow, lat- 
eral margins dentate; carapace glabrous to 
faintly rugose; with median, submedian and 
lateral carinae dentate; post-cervical carina 
absent. Eye with cornea pigmented; eye- 
stalk relatively elongate. Antennal scapho- 
cerite (acicle) just reaching distal margin of 
peduncle article 4, slightly arcuate. Maxil- 
liped 3, exopod not clearly bent; ischial 
crest well developed. Pereopod 1 , chelipeds 
asymmetrical, with spines on anterior (up- 



per) margin of propodal palm and dactylus. 
See Table 1 for branchial formula. Pleopo- 
dal rami slender; pleopod 1 in female uni- 
ramous; pleopods 2-5 with appendix inter- 
na. Lateral uropodal ramus with transverse 
suture. Telson longer than wide, with dorsal 
submedian fixed spines (Table 1). 

Remarks. — The definition of this genus 
contains some uncertainties, as the type 
species, A. pilocheira Sakai, 1987, is 
known only from the holotypic female. Im- 
portant characters of pleopods 1 and 2 of 
the male are is thus unknown, although the 
absence of pleopod 1 was deduced from 
other species placed in the genus. Three of 
the species assigned to this genus by Sakai 
& de Saint Laurent (1989) and Sakai 
(1994), A. caespitosa (Squires, 1979), A. 
hirsutimana (Boesch & Smalley, 1972), and 
A. spinosissimus (Rathbun, 1906), do not fit 
the generic definition, in that all three have 
pleopod 1 of the male present, and the car- 
apace is covered with granules, both fea- 
tures more characteristic of Oxyrhynchaxius 
Parisi, 1917. The species A. spinulicauda 
(Rathbun, 1902), also included in Acan- 
thaxius Sakai & de Saint Laurent, 1989, is 
clearly a species of Calocarides Wollebaek, 
1908. The use of the post-cervical carina, 
present in some species, absent in others, 
also needs reassessment as a diagnostic 
character. 

The advanced characters of the genus 



VOLUME 109, NUMBER 1 



71 



Table 1. — Branchial formula for Acanthaxius pil- 
ocheira. r = reduced. 



Maxillipeds 



Pereopods 



Exopod 1 1 1 — — — — — 

Epipod 111 1111 — 

Podobranch — rr rrr — — 

Arthrobranch — — 2 2222 — 

Pleurobranch — — — — — — — — 

Setobranch — — 1 11111 



would appear to be the relatively slender 
and dentate rostrum, with its basal pair of 
spines in a supraorbital position; the form 
of the chelipeds of pereopod 1, and es- 
pecially that of the smaller chela with the 
slender dentate fingers being 1.5-2.0 times 
longer than the propodal palm; the presence 
of spines on the upper margins of the dac- 
tylus and propodus of pereopod 1 chelae; 
the absence of pleurobranchs; and the pres- 
ence of epipods on pereopods 1-4. 

Species: 
Acanthaxius amakusana (Miyake & Sakai, 

1967). Amakusa Island, Kyushu, Japan, 

20-40 m. 
Acanthaxius kirkmilleri, new species. Off 

Anguilla, 421^39 m. 
Acanthaxius miyazakiensis (Yokoya, 1933). 

Southern Miyazaki-ken, Japan, 137 m; 

Philippines, 136-210 m. 
Acanthaxius pilocheira (Sakai, 1987). Off 

Honshu Island, Japan, 360 m. 
Acanthaxius polyacantha (Miyake & Sakai, 

1967). East China Sea, 118 m. 
Acanthaxius polychaetes Sakai, 1994. Off 

Great Barrier Reef, Australia, 260 m. 

Acanthaxius kirkmilleri, 

new species 

Figs. 1, 2 

Material examined. — Holotype, USNM 
243492, 9 carapace length 12.0 mm, RA^, 
John Elliott Pillsbury sta P-984, off An- 
guilla, 18°26.4'N, 63°12.6'W, 421-439 m, 
brown mud bottom, 22 Jul 1969. 

Diagnosis. — Female: Carapace glabrous; 



rostrum slender, with 2 pairs of strong dor- 
sal teeth (apex missing); median carina 
starting at rostral base, with 2 teeth anterior 
to tubercle, 1 posterior; submedian carina 
with 4 teeth; lateral carina with single 
strong tooth posterior to basal rostral tooth; 
postcervical carina poorly defined, most 
marked anteriorly. Abdominal pleuron 1 
ventrally narrowed; pleuron 2 broad, ven- 
trally truncate; pleura 3-5 with tiny denticle 
on anterior margin, anteroventrally round- 
ed, posteroventrally slightly angled. Telson 
with lateral margin having single anterior 
tooth, 2 mobile posterolateral spines, pos- 
terior margin convex, with median tooth; 
dorsal surface with 2 pairs of teeth (poster- 
iormost tooth on right side doubled). 

Antennal scaphocerite (acicle) slender, 
acute, reaching to distal margin of peduncle 
article 4. Maxilliped 3, ischium with 3 teeth 
on posterior margin; merus with 5 teeth in- 
creasing in length distally on posterior mar- 
gin. Pereopod 1 , larger chela, ischium with 
4 teeth on posterior margin; merus with 7 
teeth and several smaller denticles on pos- 
terior margin, 5 strong distal teeth on an- 
terior margin; carpus with 3 strong teeth on 
anterior margin, several smaller denticles 
on lateral surface, including strong submar- 
ginal tooth; fingers subequal in length to 
propodal palm, propodus strongly setose on 
upper and lower surfaces, with 4 strong 
teeth on anterior margin, row of 12 sub- 
marginal teeth on ventrolateral surface, lat- 
eral surface of palm with many small acute 
denticles; fixed finger with cutting edge 
bearing numerous small rounded teeth; dac- 
tylus bearing 5 strong teeth on upper mar- 
gin, cutting edge bearing numerous small 
rounded teeth. Smaller chela, ischium, me- 
rus, and carpus as in larger chela; fingers 
about Vi longer than propodal palm, latter 
setose, with 4 strong teeth on upper margin, 
row of about 10 submarginal teeth ventral- 
ly, lateral surface with several small acute 
denticles, fixed finger with cutting edge 
straight, bearing numerous small serrations; 
dactyl bearing 4 strong teeth on upper mar- 
gin, cutting edge straight, bearing numerous 



72 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. L Acanthaxius kirkmilleri, new species, holotype (USNM 243492). A, Carapace in lateral view; B, 
Anterior carapace in dorsal view; C, abdomen in lateral view; D, telson and right uropod in dorsal view. Scale 
= 2 mm. 



small serrations. Pereopod 2, anterior and 
posterior margins of all articles setose; is- 
chium with single strong posterodistal 
tooth; merus with 3 strong teeth on poste- 
rior margin; carpus with single strong tooth 
at midlength of anterior margin. Pereopod 
3, merus with 3 teeth on posterior margin; 
propodus with 5 small clumps of short 
spines on posterior margin. Pereopod 4, 
merus with single small posterodistal tooth; 
dactylus setose; propodus with 4 small 
clumps of tiny spines on posterior margin. 



Pereopod 5, propodus with distolateral clus- 
ter of setae; dactylus setose. Pleopod 1 a 
single slender ramus, distally flagelliform. 
Pleopods 2-5 with slender appendix interna 
articulating at about proximal third of en- 
dopod. Uropod with lateral ramus having 2 
teeth on lateral margin, mobile spine at an- 
gle of suture, 6 spines along suture; mesial 
ramus with 2 teeth on lateral margin, 4 teeth 
on dorsal ridge including distal marginal 
tooth. 

Remarks. — The major differences be- 



VOLUME 109, NUMBER 1 



73 




Fig. 2. Acanthaxius kirkmilleri, new species, holotype (USNM 243492). A, Pereopod 1, larger cheliped; B, 
Pereopod 1, smaller cheliped; C, Pereopod 2; D, Pereopod 3; E, Pereopod 4; F, Pereopod 5; G, Pleopod 1; H, 
Pleopod 2. Scale (A-F) = 2 mm 



tween this western Atlantic species and the 
holotype of A. pilocheira (USNM 231418) 
lie in the lateral ramus of the uropod (lateral 
margin more spinose than A. kirkmilleri), 
the generally more spinose and slightly 
more robust chelipeds of pereopod 1 in A. 
pilocheira, and the more elongate telson 
(1.5 times longer than basal width) in A. 
kirkmilleri (1.25 times longer than basal 
width in A. pilocheira). 



The differences between the present spe- 
cies and the other Japanese congeners are 
easily discerned: A. miyazakiensis is a far 
more setose species, especially on the che- 
lipeds and carapace; A. polyacantha pos- 
sesses a markedly tuberculate carapace, and 
relatively more robust chelipeds; A. amak- 
usana has a rostrum shorter than the eye- 
stalks, and a relatively broader telson. The 
Australian species A. polychaetes is more 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



setose, especially on the carapace and ab- 
domen, and possesses squatter and more se- 
tose chelipeds of pereopod 1 

Etymology. — The species is named for 
Dr. Kirk D. Miller, longtime friend and 
companion. 

Acknowledgments 

I am grateful to Dr. Rafael Lemaitre for 
reading the manuscript, and making useful 
suggestions for its improvement. 

Literature Cited 

Boesch, D. F, & A. E. Smalley. 1972. A new axiid 
(Decapoda, Thalassinidea) from the northern 
Gulf of Mexico and tropical Atlantic. — Bulletin 
of Marine Science 22(l):45-52. 

Miyake, S., & K. Sakai. 1967. Two new species of 
Axiidae (Thalassinidea, Crustacea) from the 
East China Sea. — Journal of the Faculty of Ag- 
riculture, Kyushu University 14(2): 303-3 10. 

Parisi, B. 1917. I Decapodi giapponesi del Museo di 
Milano. V. Galatheidea e Reptantia. — Atti della 
Societa Italiana di Scienze Natural! 56:1-24. 

Poore, G. C. B. 1994. A phylogeny of the families of 
Thalassinidea (Crustacea: Decapoda) with keys 
to the families and genera. — Memoirs of the 
Museum of Victoria 54:79-120. 

Rathbun, M. J. 1902. Descriptions of new decapod 
crustaceans from the west coast of North Amer- 



ica. — Proceedings of the United States National 
Museum 24(1272):885-905. 

. 1906. The Brachyura and Macrura of the Ha- 
waiian Islands. — U.S. Fish Commission Bulle- 
tin for 1903, Part 3:827-930. 

Sakai, K. 1987. Two new Thalassinidea (Crustacea: 
Decapoda) from Japan, with the biogeograph- 
ical distribution of the Japanese Thalassinid- 
ea.— Bulletin of Marine Science 41 (2): 296-308. 

. 1994. Eleven species of Australian Axiidae 

(Crustacea: Decapoda: Thalassinidea) with de- 
scriptions of one new genus and five new spe- 
cies. — The Beagle, Records of the Museums 
and Art Galleries of the Northern Territory 1 1 : 
175-202. 

, & M. de Saint Laurent. 1989. A check list 



of Axiidae (Decapoda, Crustacea, Thalassinid- 
ea, Anomura), with remarks and in addition de- 
scriptions of one new subfamily, eleven new 
genera and two new species. — Naturalists 3:1- 
104. 

Squires, H. J. 1979. Axiopsis caespitosa (Thalassinid- 
ea, Axiidae), a new species from the Pacific 
coast of Colombia. — Canadian Journal of Zo- 
ology 57(8):1584-1591. 

Wollebaek, A. 1908. Remarks on decapod crustaceans 
of the North Atlantic and the Norwegian Fiords 
(I & II). — Bergens Museums Aarbog 12:1-74. 

Yokoya, Y. 1933. On the distribution of decapod crus- 
taceans inhabiting the continental shelf around 
Japan, chiefly based upon the materials collect- 
ed by S. S. Soyo-Maru, during the year 1923- 
1930. — Journal of the College of Agriculture, 
Tokyo Imperial University 12(1): 1-226. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1 ):75-90. 1996 

A new genus for Anapagrides sensu De Saint Laurent-Dechance, 1966 

(Decapoda: Anomura: Paguridae) and descriptions of 

four new species 

Patsy A. McLaughlin and Janet Haig 

(PAMcL) Shannon Point Marine Center, Western Washington, 1900 Shannon Point Road, 

Anacortes, Washington 98221-4042, U.S.A.; 

(JH) Natural History Museum of Los Angeles County, 900 Exposition Boulevard, 

Los Angeles, California 90007, U.S.A. (deceased). 

Abstract. — Anapagrides De Saint Laurent-Dechance, 1966, as originally di- 
agnosed by its author, has recently been found not to agree with several es- 
sential characters of its type species, Eupagurus (Spiropagurus) facetus Melin, 
1939. Since Anapagrides necessarily is restricted to the taxon exemplified by 
its type species, the new genus, Laurentia, is proposed for the four undescribed 
species previously referred to the former genus. These species are now fully 
described and the systematic relationship of Anapagrides sensu stricto to the 
new genus is discussed. 



In a study of the hermit crabs of Maluku, 
Indonesia, collected during the Alpha Helix 
Expedition of 1975, to northern Australian 
and eastern Indonesian waters, Haig & Ball 
(1988) recognized and illustrated a species, 
referable to the genus Anapagrides as de- 
fined by De Saint Laurent-Dechance 
(1966). Haig & Ball (1988) declined to de- 
scribe or name their taxon as a new species 
in deference to Michele de Saint Laurent, 
who had indicated, at the time of her orig- 
inal generic diagnosis (De Saint Laurent- 
Dechance 1966) and subsequently (De 
Saint Laurent 1968), that three new 
IndoPacific species of Anapagrides re- 
mained to be described. Although De Saint 
Laurent-Dechance (1966) based her diag- 
nosis of Anapagrides on these undescribed 
taxa, she designated the nominal species 
Eupagurus (Spiropagurus) facetus Melin, 
1939, as the type of the genus. 

McLaughlin & Sandberg (1995) recently 
completed a review of three of Melin's 
(1939) species, including Anapagrides fa- 
cetus. These authors found that Melin's spe- 
cies differed from De Saint Laurent-De- 
chance's (1966) generic diagnosis to such a 



significant extent, that a major emendation 
of Anapagrides was required. Anapagrides 
sp. of Haig & Ball (1988), while agreeing 
with Anapagrides sensu De Saint Laurent, 
can no longer be assigned to Anapagrides 
as emended. In addition to Haig & Ball's 
(1988) Maluku specimen, we have now 
been able to examine De Saint Laurent's 
undescribed species. These are described 
herein as new species in Laurentia, new ge- 
nus. 

Specimens for this review are from the 
Dutch Siboga, U.S. Philippine Albatross, 
U.S., and Australian/Indonesian Alpha He- 
lix expeditions. The holotype of Haig & 
Ball's species has been deposited in the In- 
donesian National Institute of Oceanogra- 
phy, Jakarta (NIOJ). Specimens from the 
Albatross Expedition have been returned to 
the National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
(USNM), and the Siboga material to the 
Zoologisch Museum, Amsterdam (ZMA). 
One measurement, shield length (SL), mea- 
sured from the tip of the rostrum to the mid- 
point of the posterior margin of the shield, 
provides an indication of animal size. 



76 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Laurentia, new genus 

Anapagrides De Saint Laurent-Dechance, 
1966:262 (in part).— Miyake, 1978:142 
(in part), see remarks. 

Anapagrides: De Saint Laurent, 1968: 
1115.— Haig & Ball, 1988:177; not An- 
apagrides De Saint Laurent-Dechance, 
1966. 

Type species. — Laurentia albatrossae, 
new species. Gender: feminine. 

Diagnosis. — Eleven pairs of trichobran- 
chiate gills. Rostrum narrowly triangular, 
produced beyond bases of ocular acicles. 
Ocular acicles slender. Antennular pedun- 
cles with elongate ultimate segment often 
provided with 1 or more long setae at distal 
margin. Antennal peduncle with supernu- 
merary segmentation. Maxillule with exter- 
nal lobe of endopod somewhat produced, 
not recurved. Crista dentata of third max- 
illiped without accessory tooth. Stemite of 
third maxillipeds unarmed, but with shal- 
low median depression. Chelipeds unequal 
or subequal, right appreciably stouter. Am- 
bulatory legs with elongate dactyls; carpi 
with dorsodistal spine. Fourth pereopods 
with single row of scales in propodal rasp. 
Stemite of fifth pereopods developed as sin- 
gle small subovate or subquadrate lobe. 
Coxa of left fifth pereopod in males with 
moderately long or long, sometimes coiled, 
sexual tube (Fig. IE, F) provided with ter- 
minal tuft of stiff setae; right fifth coxa with 
gonopore; 3 uniramous or unequally bira- 
mous unpaired left pleopods. Females with 
paired gonopores; no paired pleopods, un- 
paired left pleopods on somites 2-5. Uro- 
pods markedly asymmetrical. Telson with 
transverse suture only weakly indicated; 
posterior lobes usually asymmetrical, ter- 
minal margins oblique; posterolateral mar- 
gins delineated at least on left. 

Etymology. — This genus is named in 
honor of the noted French carcinologist, 
Michele de Saint Laurent. 

Remarks. — Miyake (1978) included An- 
apagrides sensu lato among the hermit crab 
genera reported from Japan. Although he 



cited the gill structure as being phyllobran- 
chiate, the remainder of his generic diag- 
nosis was based on that of De Saint Lau- 
rent-Dechance (1966); A. facetus was listed 
as the only species. Miyake (1978) gave no 
indication that he had any personal knowl- 
edge of the taxon. 

Laurentia albatrossae, new species 
Figs. 1, 2, 3 A, B 

Holotype. — S (SL = 3.2 mm), Philip- 
pine Islands, Sulu Archipelago, 5.5 mi NW 
of Jolo Light, 06°09'N, 120°58'E, Albatross 
station 5141, 153 m; 15 Feb 1908; USNM 
275922. 

Paratype. — Ovigerous 2 (SL = 1.97 
mm), ? Banda, 9-36 m; ZMA Crust.: 
De.201763. 

Description. — Shield (Fig. lA) slightly 
longer than broad; anterior margin between 
rostrum and lateral projections concave; an- 
terolateral margins sloping; posterior mar- 
gin truncate; dorsal surface glabrous. Ros- 
trum triangular, well developed, reaching 
nearly half length of ocular acicles, termi- 
nating subacutely. Lateral projections well 
developed, triangular, with submarginal 
spine. Ocular peduncles (including corneae) 
approximately fg shield length; corneae 
slightly dilated. Ocular acicles subtriangu- 
lar, with submarginal spine. 

Antennular peduncles, when fully extend- 
ed, overreaching ocular peduncles by % to 
nearly entire length of ultimate segment; ul- 
timate segment with 1 or 2 long setae on dor- 
sodistal margin; penultimate segment with 1 
or 2 short setae; basal segment with statocyst 
region expanded laterally and dorsoventrally 
flattened, with distal hook-like process; dor- 
solateral margin with small distal spine. 

Antennal peduncles overreaching ocular 
peduncles by approximately V2 length of ul- 
timate segment. Fifth and fourth segments 
with few long setae. Third segment un- 
armed or with small ventrodistal spinule. 
Second segment with dorsolateral distal an- 
gle produced, terminating in acute spine 
and with small secondary spine on mesial 



VOLUME 109, NUMBER 1 



77 




Fig. 1. Laurentia albatrossae, new species. Holotype, USNM 275922: A, shield and cephalic appendages; 
B, dactyl of second right pereopod (mesial view); C, dactyl, propodus and carpus of left fourth pereopod (lateral 
view); D, anterior lobe of sternite of third pereopods; E, coxae and sternite of fifth pereopods (ventral view); F, 
coxa of left fifth pereopod and sexual tube (dorsal view); G, telson. Scales equal 2.0 mm (A, B), 1.0 mm (C, 
E-G), and 0.5 mm (D). 



margin distally; dorsomesial distal angle 
with prominent acute spine. First segment 
sometimes with spinule at dorsolateral dis- 
tal angle; small spine on ventrolateral mar- 
gin distally. Antennal acicle reaching to 



base of cornea or slightly beyond; termi- 
nating in acute spine and with long setae on 
mesial margin. Antennal flagellum with 2 
or 3 long and sometimes 1 or 2 shorter setae 
every 1 to 3 articles. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Laurentia albatrossae, new species. Holotype, USNM 275922: A, right cheliped (setae omitted); B, 
left cheliped (setae omitted); C, right second pereopod (lateral view); D, left second pereopod (lateral view); E, 
dactyl of left second pereopod (mesial view); F, left third pereopod (lateral view); G, dactyl of left third pereopod 
(mesial view). Scale equals 2.0 mm. 



VOLUME 109, NUMBER 1 



79 




Fig. 3. Chelipeds of species of Laurentia, new genus. L. albatrossae, new species: holotype, USNM 275922: 
A, left cheliped; B, right cheliped. ?L. albatrossae: i , ZMA: C, right cheliped; D, left cheliped. L. senticosa, 
new species: holotype, ZMA: E, right cheliped. Scales equal 1.0 mm (A, B); 0.5 mm (C); and 0.25 mm (D, E). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



Crista dentata with 10 to 15 regularly- 
spaced, small teeth. 

Right cheliped [Figs. 2A (setae omitted), 
3B] moderately elongate, rather slender. 
Dactyl approximately % length of palm; 
cutting edge with 1 small calcareous tooth 
proximally and 1 prominent calcareous 
tooth near mid-margin, row of very small 
calcareous teeth distally; terminating in 
very small corneous claw; dorsomesial 
margin not clearly delimited; dorsal surface 
convex, armed with numerous, scattered 
small spines or very fine spinules extending 
onto mesial face dorsally, and partially ob- 
scured by long setae; ventral surface with 
scattered long setae. Palm only slightly 
shorter than carpus; dorsomesial margin 
with row of slender, sometimes irregular, 
acute spines and long setae, dorsal surface 
convex, armed with scattered small spines 
and fine spinules practically obscured by 
long setae, dorsolateral margin with long 
setae proximally, but with small spines or 
fine spinules distally and on fixed finger; 
dorsal surface of fixed finger with scattered 
fine spinules also obscured by long setae, 
cutting edge with 1 calcareous tooth in 
proximal half, very small calcareous teeth 
distally, terminating in small weakly calci- 
fied claw. Carpus slightly longer than me- 
rus; dorsomesial margin with short, trans- 
verse rows of long setae proximally and 
more distal row of moderately slender, 
acute spines, dorsal surface with 3 very ir- 
regular rows of small spinules, decreasing 
in size laterally, dorsolateral margin not de- 
limited; mesial face with transverse rows of 
long setae, lateral and ventral surfaces also 
with transverse rows of long setae, but few- 
er in number. Merus with transverse rows 
of setae on dorsal surface, extending onto 
lateral face dorsally; ventrolateral margin 
with 1 prominent and 1 smaller spine dis- 
tally, ventromesial margin with 1 prominent 
spine at distal angle, 1 spine at midlength, 
and 1 transverse setal row and smaller spine 
proximally. Ischium with row of transverse 
setal tufts adjacent to mesial margin ven- 
trally. Coxa with prominent spine on ven- 



trolateral distal angle and second, smaller 
spine on ventromesial margin in distal half. 

Left cheliped [Figs. 2B (setae omitted), 
3A] (missing in paratype) slender, not much 
shorter than right. Dactyl approximately Va- 
longer than palm; cutting edge with row of 
very small corneous teeth, terminating in 
corneous claw; dorsal surface with irregu- 
lar, nearly double row of very small spi- 
nules in midline, partially obscured by long 
setae, dorsomesial margin rounded and 
armed with scattered tiny spinules. Palm 
about % length of carpus, dorsomesial mar- 
gin with 3 acute spines, 2 proximally sep- 
arated by broad space from medianly 
placed third spine, dorsal surface with nu- 
merous tiny spinules generally obscured by 
long setae and 3 slightly stronger spines 
proximally on weakly delimited dorsolat- 
eral margin; fixed finger with dorsal surface 
similarly armed with tiny spinules and long 
dense setae, cutting edge with row of very 
small calcareous teeth interspersed with 
corneous teeth, terminating in tiny corneous 
claw; ventral surfaces all with weakly de- 
fined transverse rows of long setae. Carpus 
approximately as long as merus; dorsome- 
sial margin with row of slender, acute 
spines, dorsolateral margin with slightly 
smaller row of spines, both partially ob- 
scured by long setae; lateral and mesial fac- 
es with transverse rows of long setae, ven- 
trolateral margin with small spine distally. 
Merus with long setae on dorsal, lateral and 
mesial surfaces; ventrolateral margin with 3 
prominent acute spines and transverse rows 
of long setae; ventromesial margin with 1 
spine near distal angle and 2 rather widely- 
spaced spines in proximal half. Ischium 
with long setae on ventral margin. 

Second and third pereopods (Figs. IB, 
2C-G) (right third missing in holotype, 
both second pereopods missing in paratype) 
generally similar from left to right. Dactyls 
slightly longer than propodi, slender, in dor- 
sal view slightly twisted, terminating in 
long, slender corneous claws; in lateral 
view curved ventrally; dorsal margins each 
with a row of stiff setae, mesial faces with 



VOLUME 109, NUMBER 1 



81 



long setae, ventral margins also with long 
setae and with 5 to 8 corneous spines in 
distal half to Vz. Propodi with 1 or 2 cor- 
neous spines on ventrodistal margin, long 
setae dorsally and ventrally. Carpi each 
with 1 spine on dorsal surface adjacent to 
dorsodistal angle (third) and 1 additional 
spine on dorsal surface proximally (sec- 
ond). Meri each with 1 spine on ventral 
margin in distal third (second) or unarmed 
(third), dorsal margins with tufts of long se- 
tae. Ischia unarmed. Fourth pereopod (Fig. 
IC) without preungual process; carpus with 
small spine at dorsodistal margin. Sternite 
of third pereopod (Fig. ID) with small, sub- 
ovate or subquadrate anterior lobe armed 
distally with 1 small spine. Sternite of fifth 
pereopods (Fig. IE) semisubovate, with 
long distal setae. 

Male pleopods unequally biramous; ex- 
opod well developed, endopod reduced. 
Telson (Fig. IG) with posterior lobes 
strongly asymmetrical, each with prominent 
spine at outer angle (largest on left) and 1 
or 2 additional spines on oblique terminal 
margin. 

Affinities. — Laurentia albatrossae most 
closely resembles L. senticosa, new species 
in having the dorsal surfaces of the chelae 
armed with numerous tiny spines; however, 
the right chela of L. albatrossae is more 
elongate and slender than that of L. senti- 
cosa. The dorsomesial margin of the palm 
of the right cheliped of L. albatrossae car- 
ries a row of moderate to strong spines that 
are lacking in L. senticosa. Other distin- 
guishing characters are pointed out under L. 
senticosa. 

Etymology. — Named for the U.S. Fish 
Commission Steamer Albatross. 

Distribution. — Philippines, Sulu Archi- 
pelago and Banda, Indonesia; 5 to 53 m. 

Remarks. — In her original report on An- 
apagrides sensu lato, De Saint Laurent- 
Dechance (1966:262) commented that a 
species collected in the Indo-Pacific by the 
Albatross and Siboga was very near Melin's 
(1939) species [Eupagurus (Spiropagurus) 
facetus] and that her genus Anapagrides 



was proposed for the former and two other 
new species from the same region. Of the 
specimens from the three Albatross stations 
initially considered by De Saint Laurent, 
only one, from station 5141, actually rep- 
resents Laurentia. One specimen, from 5"/- 
boga station 99 [Sulu Archipelago, 6°7.5'N, 
120°26'E], appears to be one of the speci- 
mens specifically referred to by De Saint 
Laurent-Dechance (1966). It is a male (SL 
= 1.79 mm; ZMA Crust.: De.201764) that 
closely resembles L. albatrossae, but is 
now almost entirely lacking calcification, 
and morphological details are difficult to in- 
terpret. Therefore, we tentatively assign this 
specimen (Fig. 3C, D) to L. albatrossae, but 
do not consider it a paratype. The female 
paratype presumably from the Siboga Ex- 
pedition carries only the label "? Banda, 5- 
20 fms" in what appears to be De Saint 
Laurent's hand writing. 

Laurentia sibogae, new species 
Fig. 4 

Holotype. — 9, ovigerous (SL = 1.5 
mm), Sailus Ketjil, Paternoster Island, "Si- 
boga" station 37, dredged to 18 m, 30, 31 
Mar 1899, ZMA Crust.: De.201762. 

Description. — Shield (Fig. 4A) slightly 
longer than broad; anterior margin between 
rostrum and lateral projections distinctly 
concave; anterolateral margins terraced; 
posterior margin roundly truncate; dorsal 
surface glabrous. Rostrum well developed, 
appreciably overreaching lateral projec- 
tions, triangular, terminating rather bluntly. 
Lateral projections prominently developed, 
obtusely triangular, with strong marginal or 
submarginal spine. 

Ocular peduncles (including corneae) ap- 
proximately % length of shield, moderately 
stout, corneae only slightly dilated. Ocular 
acicles narrowly triangular, terminating 
acutely and with small submarginal spine. 

Antennular peduncles elongate, when 
fully extended, overreaching distal margin 
of cornea by approximately % length of ul- 
timate segment. Ultimate segment with few 



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Fig. 4. Laurentia sibogae, new species. Holotype, ZMA Crust.: De.201762: A, shield and cephalic append- 
ages; B, right cheliped; C, left cheliped; D, right second pereopod (lateral view); E, dactyl of right second 
pereopod (mesial view); F, left third pereopod (lateral view); G, dactyl of left third pereopod (mesial view); H, 
anterior lobe of sternite of third pereopods; I, coxae and stemite of fifth pereopods; J, telson. Scales equal LO 
mm (A-G) and 0.5 mm (H-J). 



VOLUME 109, NUMBER 1 



83 



short setae on ventral surface. Penultimate 
segment glabrous. Basal segment with 1 
small acute spine- on dorsolateral margin 
distally. 

Antennal peduncles overreaching ocular 
peduncles by Vz to V2 length of ultimate seg- 
ment. Fifth and fourth segments with scat- 
tered, moderately long setae. Third segment 
with very small spinule on ventrodistal 
margin. Second segment with dorsolateral 
distal angle produced, terminating in simple 
or bifid spine; dorsomesial distal angle with 
moderately strong spine. First segment with 

1 acute spine ventrolateral margin distally. 
Antennal acicle moderately short, not 
reaching beyond proximal margin of ulti- 
mate peduncular segment. Antennal flagel- 
lum with 1 or 2 quite long and 1^ shorter 
setae practically every article. 

Third maxilliped with 7 or 8 small teeth 
on crista dentata; proximal 3 or 4 more 
prominent. 

Right cheliped (Fig. 4B) with chela mod- 
erately slender. Dactyl only slightly longer 
than palm; dorsomesial margin with 3 low 
spinulose protuberances, dorsal surface 
with few scattered very small spinules and 
sparse tufts of long setae; cutting edge with 

2 widely spaced large calcareous teeth in 
proximal % and few small calcareous teeth 
interspersed with corneous teeth distally. 
Palm with 4 widely-spaced spines or spi- 
nulose tubercles on dorsomesial margin, 
dorsal surface convex, apparently complete- 
ly unarmed, but with scattered long setae, 
dorsolateral margin not delimited on palm, 
but marked by tufts of long, stiff setae on 
fixed finger, dorsal surface of fixed finger 
also with numerous low protuberances and 
tufts of long, stiff setae; cutting edge with 
2 large and several smaller calcareous teeth; 
ventral surfaces of dactyl, palm, and fixed 
finger all with tufts of long setae. Carpus 
approximately as long as palm; dorsomesial 
margin with 2 widely spaced, strong spines 
and proximal transverse row of long stiff 
setae, dorsal surface with few transverse 
rows of long, stiff setae, dorsolateral margin 
not delimited, lateral and ventral surfaces 



with scattered long setae, 1 acute spinule at 
ventrolateral distal angle. Merus approxi- 
mately as long as carpus; dorsodistal mar- 
gin with long setae, and few scattered long 
setae on dorsal surface; ventrolateral mar- 
gin with 2 spines in distal half, ventrome- 
sial margin with small spine at distal angle. 
Ischium unarmed, but with few long setae. 

Left cheliped (Fig. 4C) with moderately 
long slender chela; dactyl and fixed finger 
lacking distinct hiatus, and not noticeably 
curved ventrally. Dactyl slightly longer 
than palm; dorsal and ventral surfaces un- 
armed, but with numerous tufts of long se- 
tae. Palm somewhat elevated in midline, 
perhaps minutely serrate and with long se- 
tae, dorsomesial and dorsolateral faces slop- 
ing, unarmed but with scattered long setae. 
Carpus approximately Vs longer than palm, 
covered dorsally, mesially and to lesser ex- 
tent laterally and ventrally with long setae; 
dorsomesial distal angle with acute spine, 
dorsolateral margin with row of 3 acute, 
rather widely-spaced spines and long setae; 
ventrolateral distal angle with acute spine. 
Merus approximately equaling length of 
carpus; dorsal and ventral surfaces with 
scattered long setae; ventrolateral margin 
with 2 acute spines distally, ventromesial 
margin with 1 acute spine at distal angle. 
Ischium with long setae on ventral margin. 

Ambulatory legs (Fig. 4D-G) generally 
similar from left to right; dactyls slender, V4 
to V3 longer than propodi; terminating in 
long slender corneous claws; dorsal mar- 
gins each with row of long, corneous, 
spine-like bristles in distal % and long setae, 
ventral margins each with row of corneous 
spines (6 to 8) in distal % or slightly more 
and additional long setae; mesial and lateral 
faces with scattered setae. Propodi with 1 
corneous spinule at least on left second and 
third; dorsal and ventral surfaces all with 
tufts of long setae. Carpi each with small 
almost transparent spine at dorsodistal mar- 
gin, few long setae on dorsal and ventral 
margins. Men with long setae dorsally and 
ventrally. Ischia also with long setae on 
ventral margins. Fourth pereopod without 



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preungual process at base of claw; carpus 
without dorsodistal spine. Stemite of third 
pereopods (Fig. 4H) with anterior lobe 
roundly subquadrate, anterior margin with 
long setae and 1 spine. Sternite of fifth per- 
eopods (Fig. 41) subovate, with median tuft 
of long setae. 

Telson (Fig. 4J) with transverse suture in- 
dicated in part by tuft of setae (left side); 
terminal margins oblique, each with 4 
spines, stronger on left and/or at outer an- 
gles. 

Affinities. — Of the four species of Lau- 
rentia currently recognized, L. sibogae ap- 
pears most closely related to L. balli new 
species. In both species the dorsomesial 
margin of the right chela is similarly armed 
with a row of spines; however, the dactyl 
of the right chela has only a few low pro- 
tuberances in L. sibogae, whereas in L. balli 
this margin is provided with a row of 
spines; the dorsal surfaces of the palm and 
fixed finger have a few small spines. Ad- 
ditionally, the dorsodistal margin of the car- 
pus of the right cheliped is unarmed in L. 
sibogae, but carries a prominent median 
spine in L. balli. 

Etymology. — Named for the Dutch re- 
search vessel "Siboga." 

Distribution. — At present, known only 
from the type locality; 18 m. 

Laurentia balli, new species 
Fig. 5 

Anapaghdes sp.: Haig & Ball, 1988:177, 
Fig. 8. 

Holotype.—S (SL = 2.0 mm), off Pulau, 
Saparua, Indonesia, 3°37.9'S, 128°38.6- 
39 'E, Alpha Helix Saparua station 3, 29 
Mar 1975, coll. E. Ball, NIOJ. 

Description. — Shield (Fig. 5A) approxi- 
mately as long as broad; anterior margin be- 
tween rostrum and lateral projections con- 
cave; anterolateral margins sloping; poste- 
rior margin roundly truncate. Rostrum tri- 
angular, produced beyond bases of ocular 
acicles, terminating subacutely. Lateral pro- 
jections well developed, triangular, termi- 



nating acutely and with marginal or sub- 
marginal spinule. 

Ocular peduncles approximately equaling 
length of shield, cylindrical, with comeae 
slightly dilated. Ocular acicles narrowly tri- 
angular, with small marginal or submarginal 
spine. 

Antennular peduncles (when extended) 
overreaching comeae by Vi to % length of 
ultimate segment. Ultimate segment with 1 
or 2 long setae on dorsodistal margin. Pen- 
ultimate segment with few scattered setae. 
Basal segment with statocyst area broad 
and dorsoventrally flattened, with acute, 
curved spine on lateral margin. 

Antennal peduncles equaling or slightly 
overreaching distal margins of comeae. 
Fifth and fourth segments with few scat- 
tered setae. Third segment with acute spine 
on ventral margin. Second segment with 
dorsolateral distal angle produced, termi- 
nating in acute spine, lateral margin with 
few setae; dorsomesial distal angle with 
moderately strong spine, mesial and lateral 
surfaces with few long setae. First segment 
with spinule on dorsolateral distal angle and 
1 acute spine ventrolaterally. Antennal aci- 
cle reaching beyond base of fifth peduncu- 
lar segment, slightly arcuate, terminating in 
small spine and 1 to 3 long setae. Antennal 
flagellum moderately short, with 1 or 2 long 
and to 4 short setae every article. 

Crista dentata of third maxilliped with 10 
to 13 small teeth, most proximal 3 largest. 

Right cheliped with dactyl slightly short- 
er than palm; dorsomesial margin with row 
of very small, moderately widely-spaced 
spinules, surfaces all with numerous long 
setae; cutting edge with 2 prominent cal- 
careous teeth in proximal half, row of small 
calcareous teeth distally; terminating in 
small corneous claw. Palm (Fig. 5B) slight- 
ly shorter than carpus; dorsomesial margin 
with row of 4 slender spines, dorsolateral 
margin not delimited, dorsal surface with 
scattered setae and few spinules laterally, 
more numerous on fixed finger; lateral face 
with short transverse rows of long setae; 
cutting edge of fixed finger with 2 promi- 



VOLUME 109, NUMBER 1 



85 




Fig. 5. Laurentia balli, new species. Holotype, NIOJ: A, shield and cephalic appendages; B, chela of right 
cheliped; C, chela of left cheliped; D, carpus of second right pereopod (mesial view); E, anterior lobe of sternite 
of third pereopods. Scale equals 1.0 mm (A-D) and 0.5 mm (E). 



nent and 2 smaller calcareous teeth proxi- 
mally, row of small calcareous teeth distal- 
ly. Carpus slightly longer than merus; dor- 
somesial margin with 2 widely-spaced 
acute spines distally and 2 spinulose pro- 
tuberances proximally, 1 additional spine 



on distal margin medially; ventrolateral 
margin with acute spine distally; all sur- 
faces with scattered tufts of long setae. Me- 
rus subtriangular; dorsal margin with tufts 
of moderately long setae; ventrolateral dis- 
tal angle with acute spine and tuft of long 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



setae. Ischium with scattered long setae 
dorsally and ventrally. 

Left chela (Fig. 5C) with distinct hiatus 
between dactyl and fixed finger. Dactyl 
slightly longer than palm; unarmed but with 
long setae on mesial face; cutting edge with 
row of small corneous teeth, terminating in 
small corneous claw. Palm with dorsal sur- 
face slightly elevated in midline, with few 
scattered moderately short setae mesially 
and laterally, dorsolateral margin with long 
setae, more numerous on fixed finger; cut- 
ting edge of fixed finger with row of small 
calcareous teeth. Carpus slightly longer 
than palm, approximately equaling length 
of merus; dorsolateral margin with row of 
3 widely-spaced spines, dorsomesial margin 
with 1 strong spine distally and 3 short, 
transverse ridges and long setae in proximal 
y-i, extending onto mesial face dorsally; lat- 
eral face with scattered somewhat shorter 
setae. Merus subtriangular; dorsal margin 
with row of moderately long setae; ventro- 
mesial margin with long setae and 1 acute 
spine at distal angle; ventrolateral margin 
row of long setae and 2 prominent spines 
in distal half. Ischium damaged in holotype. 

Ambulatory legs with dactyls at least Wi 
times longer than propodi; in lateral view, 
straight (second left missing in holotype) or 
curved (third); dorsal surfaces each with 
row of long, moderately stiff setae; mesial 
and lateral faces each with few scattered se- 
tae; ventral margins each with 6 corneous 
spines in distal %. Propodi with scattered 
setae dorsally and ventrally; ventrodistal 
margins each with corneous spine. Carpi 
each with small spine on dorsal surface near 
distal angle, second (Fig. 5D) with addi- 
tional small spine in proximal half, long se- 
tae dorsally and to lesser extent ventrally. 
Meri and ischia with long setae dorsally and 
ventrally. Fourth pereopod with unarmed 
carpus. Sternite of third pereopods with an- 
terior lobe (Fig. 5E) roundly subquadrate, 
with 4 small marginal spines. Sternite of 
fifth pereopods subquadrate, with long mar- 
ginal setae. 

Telson (now missing from holotype) with 



transverse suture not distinct; posterior 
lobes markedly asymmetrical, with terminal 
margins oblique, each with 3 or 4 spines, 
strongest at outer angle, lateral margin of 
left with narrow corneous plate. 

Affinities. — As previously mentioned, L. 
balli appears most closely allied to L. si- 
bogae. In addition to the characters already 
cited that separate the two species, there are 
differences in the armature of the anterior 
lobe of the sternite of the third pereopods 
and the asymmetry of the telsonal lobes. In 
L. balli, the anterior lobe of the sternite car- 
ries 4 marginal spines; only 1 is present in 
L. sibogae. The posterior lobes of the telson 
are strongly asymmetrical in L. balli, but 
very weakly so in L. sibogae. 

Etymology. — It gives us great pleasure to 
name this species for Eldon E. Ball, in rec- 
ognition of his work on Pacific hermit 
crabs. 

Distribution. — Saparua and possibly 
Banda, Indonesia. 

Remarks. — The figures of ''Anapagrides 
sp." (Haig & Ball 1988) are generally di- 
agrammatic, and some do not accurately de- 
pict certain characters. Specifically, the dor- 
solateral distal angle of the second antennal 
peduncular segment is illustrated as a sim- 
ple spine; however, there actually is a sec- 
ond spine on the mesiodistal margin; the 
first peduncular segment has a small spinule 
at the dorsolateral distal angle. The dactyl, 
fixed finger, and dorsolateral surface of the 
palm of the right cheliped appear unarmed 
in Haig & Ball's Fig. 8B. As may be seen 
in our figure (Fig. 5B), the dactyl has a few 
small spines on the dorsomesial margin and 
there are a few small spines laterally on the 
palm; on the fixed finger, a few spinules are 
present on the dorsal surface proximally 
and on the dorsolateral margin. The right 
second pereopod appears to have only a 
dorsodistal spine on the carpus in Haig & 
Ball's fig. 8E. In lateral view, the typical 
second spine on the proximal half of the 
dorsal surface is not readily observable; 
however, in mesial view (Fig. 5D) it is quite 
obvious. 



VOLUME 109, NUMBER 1 



87 



One S (SL = 2.32 mm, ZMA Crust.: 
De. 201 765) lacking chelipeds and ambula- 
tory legs, from "Siboga" station "Banda," 
is tentatively assigned to L. balli. It shares 
with the holotype the strongly asymmetrical 
telsonal lobes, and similarly armed anterior 
lobe of the stemite of the third pereopods. 
However, as variation among species of 
Laurentia is not known, we are not consid- 
ering this Banda specimen a paratype. 

Laurentia senticosa, new species 
Figs. 3E, 6 

Holotype. — S (SL = 1.79 mm), Seram 
Ceram, Indonesia, 02°28.5'S, 131°3.3'E, 5/- 
boga station 166, 118 m, 22 Aug. 1899, 
ZMA Crust.: De.201760a. 

Paratypes.—l S, I 9 (SL = 1.28, 1.40 
mm), Seram Ceram, Indonesia, 02°28.5'S, 
131°3.3'E, Siboga station 166, 118 m, 22 
Aug 1899, ZMA Crust.: De.20 1760b. 

Description. — Shield (Fig. 6A) slightly 
to considerably longer than broad; anterior 
margin between rostrum and lateral projec- 
tions concave; anterolateral margins sloping 
or terraced; posterior margin truncate. Ros- 
trum triangular, well developed, reaching 
approximately half length of ocular acicles, 
terminating bluntly or subacutely. Lateral 
projections well developed, triangular, with 
marginal or submarginal spine. 

Ocular peduncles subcylindrical, approx- 
imately % shield length; corneae dilated. 
Ocular acicles subtriangular, with small 
submarginal spine. 

Antennular peduncles overreaching ocu- 
lar peduncles by about % length of ultimate 
segment; ultimate segment with 1 or 2 long, 
simple or plumose setae on dorsodistal mar- 
gin; penultimate segment glabrous; basal 
segment with statocyst region expanded lat- 
erally and dorsoventrally flattened, dorso- 
lateral margin with long slender spine dis- 
tally. 

Antennal peduncles overreaching ocular 
peduncles by %-% length of ultimate seg- 
ment. Fifth and fourth segments with few 
long setae. Third segment unarmed or with 



small ventrodistal spinule. Second segment 
with dorsolateral distal angle produced, ter- 
minating in acute spine and sometimes with 
prominent slender spine. First segment with 
small spine on ventrolateral margin distally. 
Antennal acicle reaching to base of cornea 
or slightly beyond, terminating in acute 
spine and with long setae on mesial margin. 
Antennal flagellum missing. 

Crista dentata with 5-10 large or irreg- 
ularly-sized teeth, largest 3 or 4 often prox- 
imal. 

Right cheliped (Figs. 3E, 6B) moderately 
long and stout; with hiatus between dactyl 
and fixed finger. Dactyl approximately Vs 
length of palm; cutting edge with 2 widely- 
spaced calcareous teeth in proximal %, row 
of very small calcareous teeth distally; ter- 
minating in very small corneous claw; dor- 
somesial margin with row of very small 
spines; dorsal surface convex, with scat- 
tered long setae, midline with irregular 
rows of very small spinules; ventral and 
mesial surfaces also with scattered long se- 
tae. Palm as long or slightly longer than 
carpus; dorsomesial margin not delimited, 
dorsal surface convex, armed with scat- 
tered, very fine spinules extending onto me- 
sial face and few scattered long setae, dor- 
solateral margin with few long setae but 
otherwise not delimited proximally, with 
fine spinules distally and on fixed finger; 
dorsal surface of fixed finger with scattered 
fine spinules and long setae; cutting edge 
with 3 broad calcareous teeth, terminating 
in small corneous claw. Carpus approxi- 
mately equal to length of merus; dorsome- 
sial margin with row of moderately slender, 
acute spines, dorsal surface with 2 to 4 ir- 
regular longitudinal rows of small spinules, 
dorsolateral margin not delimited; mesial 
face with transverse rows of long setae, lat- 
eral and ventral surfaces also with scattered 
long setae. Merus with transverse rows of 
setae on dorsal margin, extending onto lat- 
eral face dorsally; ventrolateral margin with 
2 prominent spines distally, ventromesial 
margin with 1 prominent spine at distal an- 
gle, 1 additional spine originating from se- 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 6. Laurentia senticosa, new species. Holotype, ZMA Crust.: De.20 1760a: A, shield and cephalic ap- 
pendages; B, right cheliped; C, left cheliped; D, right second pereopod (lateral view); E, left third pereopod 
(lateral view); F, anterior lobe of sternite of third pereopods; G, telson. Scale equals 1.0 mm (A-E) and 0.5 mm 
(F, G). 



VOLUME 109, NUMBER 1 



89 



tal protuberance at midlength, and 1 trans- 
verse setal row and smaller spine proxi- 
mally. Ischium with setae mesially and ven- 
trally. Coxa sometimes with prominent 
spine on ventrolateral distal angle. 

Left cheliped (Fig. 6C) slender; not ap- 
preciably shorter than right but much less 
robust. Dactyl approximately equal to or 
slightly longer than palm; cutting edge with 
row of very small corneous teeth, terminat- 
ing in corneous claw; dorsal surface con- 
vex, with 4 or 5 spinules and tufts of long 
setae in midline, low protuberances and 
long setae on dorsomesial margin. Palm 
about 2/3 length of carpus; dorsal surface 
convex, armed with irregular longitudinal 
rows of tiny spinules, fewer on dorsomesial 
side and on fixed finger, dorsolateral margin 
with row of slender acute spines; palm and 
fixed finger both with scattered long setae; 
cutting edge of fixed finger with row of 
widely-spaced corneous teeth, terminating 
in small corneous claw; ventral surfaces all 
with weakly defined short transverse rows 
of long setae. Carpus approximately as long 
as merus; dorsolateral margin with row of 
slender, acute spines, dorsomesial margin 
also with row of spines strongest distally, 
both rows partially obscured by long setae; 
mesial and ventral faces with transverse 
rows of long setae; lateral face with scat- 
tered long setae, ventrolateral margin with 
acute spine distally. Merus with long setae 
on dorsal, lateral and ventral surfaces; ven- 
trolateral margin with 2 prominent acute 
spines distally, continued as row of smaller 
spinules on lateral face ventrally; ventro- 
mesial margin with 1 spine near distal an- 
gle. Ischium with long setae on ventral mar- 
gin. Coxa with prominent spine at ventro- 
lateral distal angle. 

Second and third pereopods (Fig. 6D, E) 
(holotype with only third left remaining) 
generally similar from left to right. Dactyls 
slightly to considerably longer than propo- 
di, slender; in dorsal view, slightly twisted; 
in lateral view, curved ventrally; terminat- 
ing in long, slender corneous claws; dorsal 
margins each with a row of stiff setae, me- 



sial faces with long setae, ventral margins 
also with long setae and 5-8 long corneous 
spines in middle half to %. Propodi with 1 
or 2 corneous spines on ventrodistal mar- 
gin, long setae dorsally and ventrally. Carpi 
each with 1 spine on dorsal surface adjacent 
to dorsodistal angle (third) and 1 additional 
spine on dorsal surface proximally (sec- 
ond). Meri each with 1 spine on ventral 
margin in distal third (second) or unarmed 
(third), dorsal margins with tufts of long se- 
tae. Ischia unarmed. Fourth pereopod with 
small preungual process at base of claw; 
carpus unarmed. Sternite of third pereopods 
with small, subovate or subtriangular ante- 
rior lobe (Fig. 6F) usually with 1 or 2 mar- 
ginal spines. 

Telson (Fig. 6G) with posterior lobes 
only slightly asymmetrical, each outer an- 
gle acutely developed and 1 or 2 slight pro- 
tuberances and tufts of setae on oblique ter- 
minal margins. 

Affinities. — In having the dorsal surfaces 
of the palms of both chelipeds armed with 
spinules, L. senticosa closely resembles L. 
albatrossae. However, the two species are 
immediately distinguished by the presence 
in the latter species of a row of distinct 
spines on the dorsomesial margin of the 
palm of the right chela and the more nu- 
merous, smaller and more regular teeth of 
the crista dentata. The telsons of the two 
species are also distinctly different. 

Etymology. — From the Latin senticosus 
meaning full of thorns, and referring to the 
spinulose dorsal surfaces of the chelae of 
this species. 

Distribution. — Currently known only 
from the type locality, Seram Ceram, In- 
donesia. 

Discussion 

As previously indicated, Melin's (1939) 
Eupagurus (Spiropagurus) facetus [also cit- 
ed by Melin as Eupagurus (Catapagurus) 
facetus], not only was originally designated 
as the type species of Anapagrides, but, as 
the only nominal species assigned to the ge- 



90 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



nus, must be considered the type species 
whether or not it agrees with the generic 
diagnosis given by De Saint Laurent-De- 
chance (1966) (L. B. Holthuis, pers. 
comm.)- As emended by McLaughlin & 
Sandberg (1995), Anapagrides sensu stricto 
is characterized as follows: 1) phyllobran- 
chiate gills; 2) males with short sexual tube 
on right fifth coxa; 3) prominent accessory 
tooth on the crista dentata; 4) females with 
single gonopore on coxa of left third pereo- 
pod; 5) coxa of the male third left pereopod 
frequently with female-type gonopore. In 
contrast, Laurentia has: 1) trichobranchiate 
gills; 2) males with long sexual tube arising 
from the coxa of left fifth pereopod and 
provided with a terminal tuft of stiff setae; 

3) no accessory tooth on the crista dentata; 

4) females with paired gonopores; 5) males 
without female-type genital pore on coxa of 
left third pereopod. 

All of the species presently recognized in 
Laurentia are morphologically quite simi- 
lar. Among the materials reviewed initially 
by M. de Saint Laurent, is an additional lot 
from the Siboga Expedition, which is as- 
signable, at least in part to Laurentia; how- 
ever, none of the specimens are in suffi- 
ciently good condition now to permit us to 
assign them with confidence to any of the 
described taxa. Siboga station 260 contains 
the bodies of at least five specimens of Lau- 
rentia, but also representatives of at least 
two other genera. Of the remaining append- 
ages, some most probably belong to the 
Laurentia specimens, but none can be 
matched with certitude to any of the bodies. 



Acknowledgements 

We are deeply indebted to Michele de 
Saint Laurent for having made the Alba- 
tross and Siboga specimens available for 
study. The photographs are the work of E. 
J. McGeorge. This is a scientific contribu- 
tion from the Shannon Point Marine Center. 

Literature Cited 

Haig, J., & E. E. Ball. 1988. Hermit crabs from north- 
ern Australian and eastern Indonesian waters 
(Crustacea Decapoda: Anomura: Paguroidea) 
collected during the 1975 Alpha Helix Expedi- 
tion. — Records of the Australian Museum 
(1988)40:151-196. 

McLaughlin, P. A., & L. Sandberg. 1995. Redescrip- 
tions of Gustaf Melin's 1939 "Eupagurus (Pa- 
gurillusY' exiguus, "Eupagurus (Catapagu- 
rus)" vallatus, and "Eupagurus (Spiropagu- 
rus)" facetus (Decapoda: Anomura: Paguridae) 
based on the type material. — Journal of Crus- 
tacean Biology, 15(3):569-587. 

Melin, G. 1939. Paguriden und Galatheiden von Prof. 
Dr Sixten Bocks Expedition nach den Bonin- 
Inseln 1914. — Kungliga Svenska Vetenskapsak- 
ademiens Handlingar (3)18(2):1-119. 

Miyake, S. 1978. The crustacean Anomura of Sagami 
Bay: 1-200 (English), 1-161 (Japanese). To- 
kyo: Hoikusha Publishing Co. 

Saint Laurent-Dechance, M. de. 1966. Remarques sur 
la classification de la famille des Paguridae et 
sur la position systematique d' Iridopagurus de 
Saint Laurent. Diagnose d' Anapagrides gen. 
nov. — Bulletin du Museum National d'Histoire 
Naturelle (2) 38(3):257-265. 

Saint Laurent, M. de. 1968. Revision des genres Ca- 
tapaguroides et Cestopagurus et description de 
quatre genres nouveaux. I. Catapaguroides A. 
Milne Edwards et Bouvier et Decaphyllus nov. 
gen. (Cmstaces Decapodes Paguridae) (suite). — 
Bulletin du Museum National d'Histoire Natu- 
relle (2) 39(6) (1967): 1100-1 119. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(l):91-96. 1996 

Rediscovery of Cymodocella algoense from South Africa 
(Crustacea: Isopoda: Sphaeromatidae) 

Brian Kensley and Mary Bursey 

(BK) Department of Invertebrate Zoology, National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 20560, U.S.A.; 
(MB) East London Museum, P.O. Box 11021, Southernwood 5213, South Africa 

Abstract. — The tiny sphaeromatid isopod Cymodocella algoense, which is 
characterized by a bulbous tubular posterior pleotelson, is rediscovered 120 
years after having been described from a single specimen from Algoa Bay. 
The new material was collected from the delta of the Nahoon River on the east 
coast of South Africa. 



Stebbing (1875) described a single tiny 
sphaeromatid isopod taken from the sand 
and fragments in a collection of sponges 
and gorgonians sent to him from Algoa 
Bay, South Africa. The species has not been 
recorded since. In 1910, Stebbing cast some 
doubt on the accuracy of the original local- 
ity data. Barnard (1940), in his list of Cy- 
modocella species from South Africa, men- 
tions C algoensis with the annotation "spe- 
cies dubia, juv." thereby casting uncertainty 
on its identity. 

In the course of an ecological survey of 
the benthos of the flood-tidal delta of the 
Nahoon River estuary, East London, on the 
east coast of South Africa, seven intertidal 
and seven subtidal sites were sampled by 
portable hydraulic suction sampler. Seven 
tiny adult female sphaeromatids were col- 
lected at two of the sites. On examination 
they proved to be the long forgotten Cy- 
modocella algoense, which is here rede- 
scribed. The Nahoon River estuary is about 
200 km north-east of the type locality, Al- 
goa Bay. Three specimens have been de- 
posited in the National Museum of Natural 
History, Smithsonian Institution (USNM 
253240), two in the South African Muse- 
um, Cape Town, and two in the East Lon- 
don Museum, South Africa. 



Family Sphaeromatidae 

Subfamily Dynameninae 

Cymodocella algoense (Stebbing, 1875) 

Figs. 1-3 

Sphaeroma algoense Stebbing, 1875:187, 
pi. 15, fig. 3. 

Cymodocella algoensis. — Stebbing, 1905: 
30; 1910:430.— Barnard, 1940:492 (spe- 
cies dubia). — Bruce, 1995:14. 

Material. — 5 non-ovigerous ?, total 
length 2.9-4.0 mm, dissected ? 3.3 mm; 
Nahoon River estuary flood-tidal delta, East 
London, South Africa, 1.6 m at bottom of 
channel at mouth, low spring tide, coarse 
sand, coll. M. Bursey, 22 Feb 1993.— 2 
non-ovigerous $, total length 3.0^.0 mm; 
Nahoon River estuary flood-tidal delta. East 
London, South Africa, 2 m on steep sloping 
bank 500 m from mouth, low spring tide, 
sand, coll. M. Bursey, 8 Feb 1993. 

Description. — Female: Body dorsally 
strongly arched; surface of cephalon, per- 
eon, and pleon smooth. Epistome roughly 
triangular, distally rounded, lateral margins 
slightly convex, leading to short, rounded, 
diverging 'legs'. Pleotelson reaching well 
beyond uropodal rami, consisting of broad, 
inflated basal half having 2 obscure, sub- 
median protuberances, posterior half tubu- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Cymodocella algoense: A, Female in dorsal view, scale = 1 mm; B, Posterior pleon in ventral view; 
C, Female in lateral view; D, Epistome; E, Antennule; F, Antenna; G, Mandible; H, Grinding surface of man- 
dibular molar; I, Mandibular palp; J, Maxilliped; K, Maxilla 1; L, Maxilla 2. 



VOLUME 109, NUMBER 1 



93 




Fig. 2. Cymodocella algoense: A-G, Pereopods 1-7 respectively. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Cymodocella algoense: A-E, Pleopods 1-5 respectively. 



lar, apically slightly narrowed with circular 
foramen. 

Antennular peduncle with robust basal 
article twice length of article 2, flanking 
and reaching anteriorly beyond epistome; 
article 3 slightly shorter and about half 
width of article 2; flagellum of 9 articles, 
single aesthetasc present on articles 3-8. 
Antennal peduncle of 5 articles, articles 1 
and 2 short, subequal; articles 3-5 increas- 
ing in length distally; flagellum of 1 1 arti- 
cles. Mandible with sclerotized incisor hav- 
ing 3 cusps; lacinia mobilis with 2 cusps; 2 



short plus 2 more elongate fringed spines 
in spine-row; molar truncate, grinding sur- 
face subcircular, ridged, with marginal se- 
tae; palp with 4 fringed setae distally on 
article 2, 8 stout fringed setae on article 3. 
Maxilla 1 , outer ramus with 5 broad simple 
and 4 denticulate distal spines; inner ramus 
with 4 circumplumose setae. Maxilla 2, in- 
ner ramus with about 12 mesiodistal cir- 
cumplumose setae; outer ramus with 4 
elongate denticulate spines distally on each 
lobe. Maxilliped, endite distally having 7 
stout, basally circumplumose and distally 



VOLUME 109, NUMBER 1 



95 



rounded setae, with single coupling hook 
on mesial margin; articles 2-5 of palp dis- 
tomesially setose, articles 2 and 3 mesiodis- 
tally somewhat lobed. Pereopod 1 more ro- 
bust but shorter than following pereopods; 
basis having pile of short stiff setules on 
dorsal surface; ischium with several simple 
setae on dorsal surface; merus with single 
stout anterodistal and posterodistal fringed 
seta; carpus triangular, with very short free 
anterior margin, with stout posterodistal 
fringed seta; propodus with 3 stout fringed 
setae on posterior surface; dactylus having 
bilobed accessory spine at base of unguis. 
Pereopod 2 more slender than following 
legs; basis with pile of short stiff spinules 
on anterior surface; ischium with 3 elongate 
simple setae on anterior surface; merus with 
2 simple anterodistal setae; carpus rectan- 
gular, % length of propodus. Pereopods 3- 
7 essentially similar, pereopod 7 slightly 
more slender than preceding legs; basis and 
ischium with pile of short spinules on an- 
terior surface; ischium with 4-6 elongate 
simple setae on anterior surface; merus 
much shorter than ischium, with 2-A simple 
setae on rounded anterodistal lobe; carpus 
with single bifringed seta distally, except in 
pereopod 7 which has 6 bifringed setae; an- 
terior surfaces of merus, carpus, and prop- 
odus having numerous short scales/spi- 
nules; dactylus with bifid accessory spine 
at base of unguis. Pleopods 1-3 each with 
elliptical exopod bearing plumose marginal 
setae; endopod triangular, subequal in 
length to exopod in pleopod 1, reaching 
well beyond exopod in pleopods 2 and 3, 
bearing plumose marginal setae. Pleopod 4, 
both rami pleated, exopod with single short 
distal seta. Pleopod 5, both rami pleated, 
exopod with transverse suture in distal half, 
armed with 3 spinulose lobes. Uropod with 
exopod freely articulating under endopod, 
elliptical, considerably shorter than endo- 
pod, latter distally rounded. 

Male: unknown. 

Remarks. — Bruce (1995), in a revision of 
the tube-tailed sphaeromatids, demonstrated 
that the genus Cymodocella was polyphy- 



letic, defined a second genus, Diclidocella, 
to accommodate some of the species, 
moved two southern African species to Is- 
chyromene, and removed C. hawaiiensis 
Bruce, 1994, from the group altogether. 

The genus Cymodocella Pfeffer, 1887, 
currently contains four species (see Bruce 
1995:5). The type species, C tubicauda 
Pfeffer, 1887, was redescribed by Brandt & 
Wagele (1989). Cymodocella algoensis can 
now be included in this group, agreeing 
well with the revised diagnosis provided by 
Bruce (1995). Although Stebbing's descrip- 
tion of C algoense is limited, the figure of 
the whole animal in dorsal view shows the 
distinctively shaped convex-sided pleotel- 
sonic tube. The remaining four South Af- 
rican species are of uncertain generic status, 
and require redescription, given the lack of 
detail in existing descriptions. 

Acknowledgments 

The first author thanks Leon Engelbrecht, 
Robson Ntsabo, and Deon Smit for helping 
to operate the hydraulic pump during sam- 
pling. Dr. Niel Bruce of the Zoological Mu- 
seum, Copenhagen, commented on the 
manuscript, and generously shared his un- 
published work on the tube-tailed sphaero- 
matids, which greatly improved this short 
contribution. We are also grateful to Ms. 
Marilyn Schotte for reading a draft of the 
paper. 

Literature Cited 

Barnard, K. H. 1940. Contributions to the crustacean 
fauna of South Africa. XII. Further additions to 
the Tanaidacea, Isopoda, and Amphipoda, to- 
gether with keys for the identification of the 
hitherto recorded marine and fresh-water spe- 
cies. — Annals of the South African Museum 
32(5):381-543. 

Brandt, A., & J.-W. Wagele. 1989. Redescriptions of 
Cymodocella tubicauda Pfeffer, 1887, and Ex- 
osphaeroma gigas (Leach, 1818) (Crustacea, Is- 
opoda, Sphaeromatidae). — Antarctic Science 1: 
205-214. 

Bruce, N. L. 1994. New records of marine isopod 
crustaceans (Sphaeromatidae) from Hawaiian 
waters. — Crustaceana 67(3):38 1—386. 



96 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



. 1995. The taxonomy and phylogeny of tube- 
tailed sphaeromatid isopods (Crustacea) with 
descriptions of new species and a new genus 
from southern Australia. — Ophelia 43:127-180. 

Harrison, K., & J. P. Ellis. 1991. The genera of the 
Sphaeromatidae (Crustacea: Isopoda): a key and 
distribution list. — Invertebrate Taxonomy 5: 
915-952. 

Pfeffer, G. 1887. Die Krebse von Siid-Georgien nach 
der Ausbeute der Deutschen Station 1882- 
1883. — Jahrbuch der Hamburgischen Wissen- 
schaftlichen Anstalten 4:43-150. 



Stebbing, T. R. R. 1875. On some new exotic sessile- 
eyed crustaceans. — Annals and Magazine of 
Natural History (4)15:184-188. 

. 1905. On the Isopoda. Pages 1-64, in W. A. 

Herdman, Report to the Government of Ceylon 
on the Pearl Oyster Fisheries of the Gulf of Ma- 
naar. Supplementary Report 23. 

. 1910. General catalogue of South African 

Crustacea. — Annals of the South African Mu- 
seum 6(4):28 1-593. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1):97-102. 1996 

Spelaeomysis villalobosi, a new species of mysidacean from 
northeastern Mexico (Crustacea: Mysidacea) 

Maria Elena Garcia-Garza, Gabino A. Rodriguez-Almaraz, and Thomas E. Bowman 

(MEG-G, GAR-A) Facultad de Ciencias Biologicas, 

Universidad Autonoma de Nuevo Leon, Ap. postal 105-F, 

Cd. Universitaria, San Nicolas de los Garza, Nuevo Leon, Mexico; 

(TEB) Department of Invertebrate Zoology, NHB-163, Smithsonian Institution, 

Washington, D.C. 20560, U.S.A. (deceased) 

Abstract. — Spelaeomysis villalobosi, a new species of mysid is described 
from subterranean habitats in Nuevo Leon, Mexico. This is the third known 
species of Spelaeomysis from Mexico. Spelaeomysis villalobosi is similar to 
Spelaeomysis quinterensis (Villalobos-Figueroa, 1951) previously described 
from caves in Tamaulipas and San Luis Potosi, but differs in the rounded shape 
of the anterolateral lobe of the eyestalk, in the 4 cusped incisor of the mandible, 
the presence of three spines on the dactylus of the maxilliped, and three claws 
on the dactylus of pereopod 1 ; the exopod of pleopod 4 is 2-segmented; the 
margin of the telson is convex. A key to the known species of Spelaeomysis 
is given. 

Resumen. — Spelaeomysis villalobosi, es una nueva especie de misidaceo hi- 
pogeo encontrada en el Estado de Nuevo Leon, Mexico. Esta especie es la 
tercera conocida de Spelaeomysis en Mexico. Spelaeomysis villalobosi presenta 
parecido morfologico con Spelaeomysis quinterensis Villalobos-Figueroa, 1951 
ubicada en cuevas de Tamaulipas y San Luis Potosi, pero existen diferencias 
notorias entre ambas especies, principalmente con respecto a la forma de los 
pedunculos oculares, la denticion del proceso incisivo de la mandil^ula, la pre- 
sencia de espinas en el dactilo del maxilipedio, el niimero de uiias en el dactilo 
del pereiopodo 1, pleopodo 4 con su exopodo con 2 segmentos en vez de 4, 
y la forma y ornamentacion del telson y el uropodo. Se incluye una clave para 
las especies de Spelaeomysis conocidas. 



The freshwater mysids of Mexico are Family Lepidomysidae Clarke, 1961 

poorly known. Antromysis cenotensis, A. Genus Spelaeomysis Caroli, 1924 

reddelli, Spelaeomysis olivae and S. quin- Spelaeomysis villalobosi, new species 

terensis have been found in hypogean hab- Figs- 1-18 

itats (Bowman 1982). Taphromysis louisi- Material examined.— WoXoiypt male, 

anae is the first freshwater epigean mysi- uANL (Universidad Autonoma de Nuevo 

dacean recorded from Mexico (Garcia-Gar- Leon) CCRMYOOl, 1 female paratype 

za et al. 1992). Spelaeomysis quinterensis (UANL CCRMY002) from a draw-well, 

was recorded from caves in Tamaulipas and Rancho Monte Carmelo, Cerralvo, Nuevo 

San Luis Potosi (Villalobos-Figueroa 1951; Leon, 100 Km Northeast of Monterrey, 

Reddell 1981), and S. olivae is known from Nuevo Leon, about 26°04'N, 99°37'W, 21 

caves in Oaxaca (Bowman, 1973), both Oct 1989, leg. Maria Elena Garcia-Garza, 1 

species have disjunct distributions. male paratype (UANL CCRMY003) from 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 1-7. Spelaeomysis villallalobosi n. sp., Male holotype: 1, Male, lateral; 2, Eyestalks; 3, Antenna 1; 4, 
Antenna 2; 5, Telson; 6, Mandible; 7, Mandible palp. 



a small spring near the Pilon River, Mon- 
temorelos, Nuevo Leon, about 25°irN, 
99°48'W, 17 May 1990, leg. M. Valdez- 
Marroquin. 3 male paratypes (UANL 
CCRMY004) from Chorrera Cave, High- 



way Linares-Galeana, Linares, Nuevo 
Leon, 24°46'N, 99°37'W, 4 Apr 1994, leg. 
Carlos Cavazos-Camacho. 

Description. — Length of holotype 7 mm. 
Caparace with triangular rostrum (Fig. 2) 



VOLUME 109, NUMBER 1 



99 




Figs. 8-11. Spelaeomysis villalobosi n. sp. Male holotype: 8, Maxilla 1; 9, Maxilla 2; 10, Maxilliped; 11, 
Pereopod 1. 



not covering eyestalks; anterolateral lobe of wider than long, produced anterolaterally 

caparace reaching to midlength of eye- into pointed lobe. 

stalks. Eyestalk without ommatidia slightly Pleuron of pereonite 7 produced poste- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Figs. 12-18. Spelaeomysis villalobosi n. sp. Male holotype: 12, Pereopod 7; 13, Pleopod 1; 14, Pleopod 2; 
15, Pleopod 3; 16, Pleopod 4; 17, Pleopod 5; 18, Uropod. 



riorly into lobe. Telson linguiform (Fig. 5), 
as long as wide at base; posterior 0.4 
armed, with 24 marginal spines; apex with 
2 long spines flanked by 2 short spines. 

Antenna 1 peduncle (Fig. 3), segment 1 
widest, about 1 . 1 times longer than segment 
2; segment 3, shortest 0.95 X as long as seg- 
ment 2; segment 1, with 4 plumose setae at 
distomedial corner and 5 naked setae at dis- 
tolateral comer; segment 2, with row of 



plumose and naked setae on medial margin 
and 2 small setae on distolateral comer; 
segment 3 with naked and plumose setae on 
medial margin and small seta on distal part 
of lateral the margin; distal lobe as wide as 
long; bearing 3 small apical setae. Flagel- 
lum with 45 segments. Antenna 2 (Fig. 4), 
scale 2.5 X longer than wide, reaching to 
first third of last segment of peduncle, fla- 
gellum broken. 



VOLUME 109, NUMBER 1 



101 



Table 1. — Differences between Spelaeomysis quinterensis and S. villalobosi. 



S. quinterensis 



S. villalobosi 



Eyestalk anterolateral lobe 

Telson length/width 

Telson shape 

Mandibular incisor 

Mandibular palp 

Mandibular palp segment 2/seg- 

ment 3 
Maxilla 2 endopod 

Maxilliped, merus of endopod 
Pereopod 1 merus 
Pereopod 1 dactyl 
Pleopod 4 exopod 
Uropod: exopod/endopod 



Rounded 

1.3X 

Narrows at anterior 

3 strong teeth 

With slightly concave margins 
1.8X 

Segment 1 longer than segment 2, 
not produced distomedially 

4 setae 
2 setae 

1 terminal claw 
4 segments 
1.25X 



Pointed 

l.OX 

Margins evenly convex 

4 strong teeth 

2.6X 

Segment 1 shorter than segment 2, 

produced distomedially 
7 setae 
6 setae 

3 terminal claws 
2 segments 
1.30X 



Left Mandible (Fig. 6), incisor with 4 
conspicuous cusps; lacinia mobilis bearing 
5 denticles; spine row with 6 naked setae; 
molar process armed with minute setae; 
palp (Fig. 7), segment 22.6 X longer than 
segment 3, both segments with stout barbed 
setae on distal part of outer margin. 

Maxilla 1 (Fig. 8), inner lobe with 4 pec- 
tinate terminal setae and 3 naked subter- 
minal setae; 3 small setae on outer margin 
and one long setae on proximal inner mar- 
gin. Outer lobe with 14 long apical setae of 
different types; inner margin with row of 
fine hairs extending onto surface. 

Maxilla 2 (Fig. 9), segment 1 shorter than 
segment 2, produced medially to midlength 
of segment 2; segment 2 with 7 setae on 
distomedial margin; exopod 3X longer than 
wide, with about 15 long plumose marginal 
setae. 

Maxilliped (Fig. 10), merus of the en- 
dopod with 7 plumose setae, carpus with 7 
plumose setae. Propodus, distal outer mar- 
gin with 2 long plumose setae and 3 smaller 
naked setae; dactylus with 3 spines and 4 
naked setae in distal part. 

Pereopod 1 (Fig. 11), endopod is robust; 
merus with 6 naked setae; outer margin 
propodus with a long setae and 3 smaller 
setae; dactylus ending in 3 claws. Pereopds 
2-7 as in Fig. 12. 

Pleopod 1 (Fig. 13), endopod with 2 long 



naked setae on medial margin and 5 long 
naked distal setae; exopod proximal seg- 
ment with 4 short and 3 longer naked me- 
dial setae; distal segment with 7 long naked 
setae. Pleopod 2 (Fig. 14), endopod elon- 
gate, 2.8 times longer than width; segment 
3 of exopod about half length of segment 
2, setae on segment 2 short. 

Pleopod 3 (Fig. 15), endopod not reach- 
ing distal part of exopod segment 1 , with 4 
small setae on inner margin; segment 2 of 
exopod with a subterminal seta. Pleopod 4 
(Fig. 16), exopod with 2 segments. Pleopod 
5 (Fig. 17), exopod subequal in length to 
endopod; segment 2 about 1.25X longer 
than segment 3, bent laterally at midlength, 
with 5 small proximolateral setae, 2 longer 
subterminal setae, and a terminal seta as 
long as 3rd segment. 

Uropod protopod (Fig. 18), medial lobe 
with numerous curved spines and a long 
tooth; proximal part with 2 slender setae. 
Endopod reaching oblique suture of exo- 
pod. Exopod, proximal segment with 15 
spines of almost equal size. 

Etymology. — Named in memory of Dr. 
Alejandro Villalobos-Figueroa, distin- 
guished Mexican carcinologist. 

Remarks. — Spelaeomysis villalobosi is 
the third species of the genus known for 
Mexico. Spelaeomysis quinterensis (Villa- 
lobos-Figueroa, 1951) and S. villalobosi, 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



new species, have similar distribution in 
northeastern Mexico; and S. olivae (Bow- 
man, 1973) is known only from Oaxaca 
state in southern Mexico. Spelaeomysis vil- 
lalobosi is similar to S. quinterensis; the 
principal differences are listed in Table 1. 
The seven known species of Spelaeomysis 
may be distinguished by the following key. 

Key to the Species of Spelaeomysis 

1. Eyes with small anteriolateral area of 
ommatidia 2 

- Eyes without visual elements 3 

2. Apical spine of telson about 2X length 

of 2 flanking spines S. serratus 

(Page, 1924) 

- Apical spine of telson more than 6X 
length of 2 flanking spine ... 5. cardisomae 

Bowman 1973 

3. Eyestalks fused medially, forming single 
plate S. longipes 

(Pillai & Mariamma, 1963) 

- Eyestalks separate 4 

4. Eyestalks produced anterolaterally into 
subtriangular lobe 5 

- Eyestalks subquadrilangular, not pro- 
duced anterolaterally 6 

5. Pleopod 4 with LAV 4 segmented exo- 
pod; telson about 1.2; lateral margins 
slightly concave near midlength 

.5. quinterensis (Villalobos-Figueroa, 1951) 

- Pleopod 4 with LAV 2 segmented exo- 
pod; telson about 1.0; lateral margins 
evenly convex S. villalobosi n. sp. 

6. Telson LAV 1.5, with about 20 marginal 
spinal 

S. nuniezi Bacescu & Orghidan, 1971 

- Telson LAV 1.1-1.2, with about 40 mar- 
ginal spines S. olivae Bowman, 1973 

Acknowledgments 

We are grateful to Dr. Elva Escobar-Bri- 
ones of the Instituto de Ciencias del Mar y 
Limnologia, UNAM, Mexico for her com- 
mentaries and criticism during the elabora- 
tion of the original draft. Besides, we sin- 



cerely thank two anonymous reviewers for 
their comments on this manuscript. Part of 
this work was carried out in Washington by 
the second author during a Short-term Vis- 
itor Award from the Smithsonian Institu- 
tion's Office of Fellowships and Grants. 

Literature Cited 

Bacescu M., & T. Orghidan. 197 L Antromysis cuban- 
ica n. sp. et Spelaeomysis nuniezi n. sp. mysis 
cavernicoles nouvelles de Cuba. — Revue Rou- 
maine de Biologie, serie de Zoologie 16:225- 
231. 

Bowman, T. E. 1973. Two new American species of 
Spelaeomysis (Crustacea: Mysidacea) from a 
Mexican cave and land crab burrows. — Asso- 
ciation for Mexican Cave Studies, Bulletin 5: 
13-20. 

. 1982. Mysidacea. Pp. 201-202 in S. H. Hurl- 

bert & A. Villalobos-Figueroa, eds.. Aquatic Bi- 
ota of Mexico, Central America and the West 
Indies. San Diego State University Foundation, 
San Diego, California, xv + 529 pp. 

Caroli, E. 1924. Su di un Misidaceo Cavernicolo 
{Spelaeomysis bottazii n. g., n. sp.) di Terra 
d'Otranto. — Atti dell Accademia Nazionale die 
Lincei Rendiconti, Classe die Scienze Fisiche 
Matematiche Naturali 33:512-513. 

Clarke, D. 1961. Proposal of a new name, Lepido- 
mysis, for the preoccupied mysidacean generic 
name Lepidops Zimmer, 1927. — Crustaceana 2: 
251-252. 

Fage, L. 1924. Sur un type nouveau de Mysidace des 
eaux souterraines de I'isle de Zanzibar. — Comp- 
tes Rendus Hebdomadaires des Seances de 
I'Academie des Sciences, Paris 178:2127-2129. 

Garcia-Garza, M. E., G. A. Rodn'guez-Almaraz, & T. 
E.Bowman. 1992. Distribution of the opossum 
shrimp Taphromysis louisianae (Mysidacea), 
including new far inland records. — Journal of 
Crustacean Biology 12:101-103. 

Pillai, N. K., & T. Mariamma. 1963. On the discovery 
of the primitive mysidacean family Lepidomy- 
sidae in India. — Current Science, Bangalore 32: 
219-220. 

Reddell, J. R. 1981. A review of cavernicole fauna of 
Mexico, Guatemala and Belize. — Bulletin of 
the Texas Memorial Museum 27:1-327. 

Villalobos-Figueroa, A. 1951. Un nuevo Misidaceo de 
las grutas de Quintero en el Estado de Tamau- 
lipas. — Anales del Instituto de Biologia, Univ- 
ersidad Nacional Autonoma de Mexico 22:191- 
218. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1):103-1U. 1996 

A new copepod species from California, U.S.A.: Hesperodiaptomus 
californiensis (Crustacea: Copepoda: Calanoida: Diaptomidae) 

Megan Scanlin and Janet W. Reid 

(MS) College Station Box 3505, The College of William and Mary, 

Williamsburg, Virginia 23186-3505, U.S.A.; 

(JWR*) Research Associate, Department of Invertebrate Zoology/MRC-163, 

National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560, U.S.A. (*Corresponding author) 

Abstract. — Hesperodiaptomus californiensis, new species (Copepoda: Cal- 
anoida: Diaptomidae) was collected from vernal pools in Lassen County, north- 
em California, U.S.A. It differs from its three most morphologically similar 
congeners, Hesperodiaptomus schefferi, Hesperodiaptomus victoriaensis and 
Hesperodiaptomus kiseri in having both leg 5 endopods of the male relatively 
long, the left lateral protrusion on the genital double somite of the female 
directed dorsally, and in other details. Co-occurring diaptomids were Hesper- 
odiaptomus novemdecimus and Leptodiaptomus tyrrelli. 



A new species of diaptomid copepod was 
collected in Lassen County, California, as 
part of a study of vernal pools (described 
by King et al. 1996). The new copepod oc- 
curred in three pools and at the time of col- 
lection was abundant in every pool. 

Wilson (1959) listed 16 species of Hes- 
perodiaptomus in North America; since 
then no new species has been recognized 
from this continent. However, the taxon 
Hesperodiaptomus arcticus (Marsh, 1920) 
s.l. may be composed of several cryptic 
species. Boileau & Hebert (1988) and Bo- 
ileau (1991) found genetic differences be- 
tween different North American popula- 
tions of H. arcticus, which were accompa- 
nied by subtle morphological differences. 
On the other hand, Stepanova (in Borutskii 
et al. 1991) rejected efforts by Streletskaya 
(1983, 1986) to split Siberian H arcticus 
into three taxa: H. koolensis Streletskaya, 
1983, H. judayi Streletskaya, 1986 and H. 
kurenkovi Streletskaya, 1986, because of 
the supposedly minor degree of morpholog- 
ical differences. 

Some other members of Hesperodiapto- 
mus are so similar morphologically that the 



females cannot be reliably assigned to a 
particular species with present knowledge. 
These possible species complexes are H. 
breweri (Wilson, 1958a), H. eiseni (Lillje- 
borg in Guerne & Richard, 1889), and H. 
arcticus; as well as H. kiseri (Kincaid, 
1953) and H. victoriaensis (Reed, 1958). 
The high degree of morphological similar- 
ity between H. kiseri and H. victoriaensis 
creates some difficulty in establishing the 
species from California as a new taxon, be- 
cause it shares several characters with both 
these species. 

Hesperodiaptomus species and other 
diaptomid copepods are characterized clas- 
sically by the features of the fifth legs of 
the males and females, the right antennule 
of the male and setation of the left anten- 
nule of the male and both antennules of the 
female, and features of the somites, partic- 
ularly the shape of the thoracic "wings" 
(expansions of the sixth thoracic somite). 
Pending eventual revision of the genus Hes- 
perodiaptomus and the possible discovery 
of additional morphological and genetic 
characteristics, we describe the new species 
using these traditional discriminators. 



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The copepod specimens were originally 
fixed in buffered formalin and transferred 
to 70% ethanol for long term storage. Field 
sampling methods were described by King 
et al. (1996). Descriptions were made from 
whole specimens in glycerin and/or lactic 
acid; dissected specimens were mounted ei- 
ther in CMC- 10 or in commercial polyvinyl 
lactophenol with a little Chlorazol Black E 
added. All measurements were made from 
specimens in glycerin. Specimens were de- 
posited in the collections of the United 
States National Museum of Natural History, 
Smithsonian Institution (USNM) and the 
Natural History Museum of Los Angeles 
County, California (LACM). 

Family Diaptomidae G. O. Sars 

Genus Hesperodiaptomus Light, 1938 

Hesperodiaptomus calif orniensis, 

new species 

Figs. 1-3 

Type material. — Holotype S, USNM 
264057; allotype 9, USNM 264058; para- 
types: 3 S S 3 9 9, each dissected on slide 
in PVL or in CMC- 10, and 464 S S 9 9 + 
copepodids, USNM 264059; 463 c? c? 9 9 
+ copepodids, LACM 95-50.1; all from 
California, Lassen County, pool #41, 
40°37'N, 121°03'W, immediately adjacent 
to west side of Route 44, 1.4 km north of 
Bogart Safety Rest Area/Ranger Station, 2 
m tow of wet meadow edge with plankton 
net, sample 215. 

Additional, non-type material. — 372 S S 
9 9 + copepodids, California, Lassen 
County, pool #42, 40°32'N, 12rO'W, 30 m 
west of Route 44, just south of railroad 
track crossing, 2 m tow of wet meadow lake 
edge with plankton net, sample 218, USNM 
264060. 894 SS 9 9, pool #42 bottom, 
sample 217, LACM 95-50.3. 41 SS 9 9, 
pool #41, sample 214, USNM 264061. 
400+ 6 6 9 9, pool #42 surface, sample 
216, LACM 95-50.4. 2S 6 6 9 9, Califor- 
nia, Lassen County, pool #43 (Long Lake), 
40°31'N, 120°59'W, 100 m west of Route 
44, about 0.36 km south of pool #41 (about 



1.5 km south of intersection with Route 
A21), 2 m tow of wet meadow lake bottom 
with plankton net, sample 220, USNM 
264062. 10 66 9 9, pool #43 (Long Lake), 
sample 219, LACM 95-50.2. All collected 
23 April 1992 by J. L. King, D. Gluesen- 
kamp and J. Tritt. Undissected specimens 
preserved in 70% ethanol. 

Co-occurring species (determined by 
JWR). — Pool #41: Hesperodiaptomus nov- 
emdecimus (Wilson, 1953), Leptodiaptomus 
tyrrelli (Poppe, 1888), Acanthocy clops car- 
olinianus (Yeatman, 1944), Diacyclops 
crassicaudis var. brachycercus (Kiefer, 
1927), Microcyclops rubellus (Lilljeborg, 
1901), Bryocamptus washingtonensis Wil- 
son, 1958b; pool #42: H. novemdecimus, L. 
tyrrelli, A. carolinianus, Acanthocyclops 
vernalis (Fischer, 1853) s.l., Diacyclops na- 
vus (Herrick, 1882); pool #43 (Long Lake): 
H. novemdecimus, L. tyrrelli, A. carolini- 
anus, B. washingtonensis. 

Male. — Length (mm) of holotype 2.21; 
of ten paratypes, mean = 2.14, median = 
2.18, range = 1.93-2.32. Pediger 5 (sixth 
thoracic somite) (Fig. la), lateral "wings" 
symmetrical, each with two sensilla. Uro- 
some (Fig. la) symmetrical except urosom- 
ites 1 and 4 slightly produced laterally and 
posteriorly. Urosomites 2-4 and caudal 
rami with pore-canals, middle dorsal pore- 
canal of urosomite 4 displaced slightly to 
left of dorsal midline. Inner sides of caudal 
rami with long hairs, rest of ramal surfaces 
with short sparse hairs. Rostral points (Fig. 
lb) short, acute. 

Antennules (Fig. la, c, d) reaching mid- 
dle of urosomite 2. Right antennule with 
two setae on segment 6; segment 8 with one 
seta and one spine; segment 10 with one 
seta and one stout spine reaching middle of 
segment 12; segment 11 with one seta and 
one stout spine, reaching distal end of seg- 
ment 13; segment 13 with one stout spine 
reaching past middle of segment 14; seg- 
ment 14 without spine; segment 15 with 
small acute process pointing distally; seg- 
ment 16 with two setae. Segment 23 (Fig. 
Id), process at distal end straight, reaching 



VOLUME 109, NUMBER 1 



105 



50 jam 




Fig. 1. Hesperodiaptomus californiensis, new species, male: a. Habitus, dorsal; b. Rostral points (indicated 
by arrows); c. Right antennule, segments 6-17; d. Right antennule, segments 23-25; e. Leg 2 endopod, segments 
1 and 2, posterior. 



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Fig. 2. Hesperodiaptomus californiensis, new species, male: a. Leg 5, posterior; b. Left leg 5 exopod, 
posterior; c, Left leg 5 exopod and endopod, anterior; d. Left leg 5 endopod, posterior; e. Right leg 5 endopod, 
anterior. Both scales = 50 jjim. 



end of last segment of antennule. Setation 
of left antennule as in female. 

Leg 2 endopod segment 2 (Fig. le) with- 
out defined Schmeil's organ, but with small 
transverse ridge, more or less developed on 
different specimens. 

Left leg 5 (Fig. 2a-d): Coxa, small pro- 
cess tipped with sensillum near outer pos- 
terodistal margin. Basis with lateral seta. 
Exopod segment 1 longer than segment 2, 
with haired pad on inner margin, hairs thick 
and as long as about half width of endopod. 
Exopod 2, tip forming stout, blunt, smooth 
lateral process; also with serrate digital pro- 



cess. Inner margin of exopod 2 with two 
pads, proximal pad haired, proximal hairs 
of this pad thinner and shorter than distal 
hairs; distal pad covered with rows of teeth 
and few tiny hairs near tip of exopod. Pos- 
terior surface of exopod 2 with longitudinal 
groove. Endopod reaching midlength of ex- 
opod 2, of one segment, narrowing at distal 
Vs. Tip of endopod narrowing in acute pro- 
cess; also with five small subterminal 
spines and group of subterminal hairs. 

Right leg (Fig. 2a, e): Coxa with small 
posterodistal process ending in sensillum. 
Basis with lateral seta, shallow longitudinal 



VOLUME 109, NUMBER 1 



107 




Fig. 3. Hesperodiaptomus califomiensis, new species, female: a. Habitus, dorsal; b. Rostral points (indicated 
by arrows); c, Antennule, segments 1-9; d, Leg 5, anterior; e. Tip of leg 5 endopod, posterior. 



groove on posterior surface, and rounded, 
unornamented protrusion on middle of in- 
ner margin; length of protrusion equal to 
width of endopod, its width equal to V2 
width of endopod. Outer margin of exopod 
1 with large, distally directed conical pro- 
cess on distolateral comer. Exopod 2 almost 
twice length of exopod 1 , with shallow lon- 
gitudinal groove along distal V2 of inner 



margin; lateral spine at distal % of exopod 
2, V2 thickness of endopod and Vi length of 
exopod 2, straight, finely denticulate. Ter- 
minal claw tapering gradually from en- 
larged base, twice length of exopods 1 and 
2 combined, with row of teeth extending 
from tip of claw almost to base. Right en- 
dopod without suture on posterior surface, 
suture visible on anterior surface where en- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



dopod narrows slightly at midlength; en- 
dopod tapering to blunt point; with 3-5 
spines near outer distal margin, and subter- 
minal group of small hairs. 

Female. — Length (mm) of allotype 2.39; 
of ten paratypes, mean = 2.20, median = 
2.22, range = 1.95-2.33. Prosome (Fig. 3a) 
symmetrical, thoracic wings symmetrical, 
each with two sensilla located on lateral and 
posterior margins. Genital double somite 
extended laterally in asymmetrical conical 
projections each tipped with sensillum, left 
projection directed dorsally, right projection 
laterally. Caudal rami haired in pattern sim- 
ilar to that of male. Rostral points (Fig. 3b) 
low, rounded. 

Antennule (Fig. 3a, c) reaching mid- 
length of genital double somite. Segments 
1-8 or 9 each with few tiny hairs irregularly 
scattered on posterior surface. Seta on seg- 
ment 1 long, reaching midlength of segment 
4. Appendages per segment as follows (Ro- 
man numerals = segment, Arabic numerals 
= number of setae, a = aesthetasc, sp = 
spine): 1(1 +a), II(3+a), 111(1 +a), IV(1), 
V(l+a), VI(1), VII(l+a), VIII(l-Hsp), 
IX(2+a), X(l), XI(2), Xn(l+sp+a), XIII(l), 
XIV(l+a), XV(1), XVI(l+a), XVII(l), 
XVIII(l), XIX(l+a), XX(1), XXI(l), 
XXn(2), XXIII(2), XXIV(5+a). 

Leg 2 (not figured) of all females ex- 
amined without trace of Schmeil's organ or 
transverse ridge on endopod 2. 

Leg 5 (Fig. 3d, e): Coxa with posterior 
lateral protrusion ending in sensillum. Basis 
with lateral seta reaching Vz of length of ex- 
opod 1. Endopod two-segmented, distal 
segment twice length of proximal segment. 
Endopod 2 with two setae at tip, tip pro- 
truding in acute point between these setae, 
and 3-5 subterminal teeth surrounding tip 
of endopod. Exopod 2 a little longer than 
exopod 1, claw with inner margin toothed. 
Exopod 2 with small articulated spine lat- 
eral to outer margin of exopod 3, reaching 
from slightly beyond middle to end of ex- 
opod 3. Exopod 3 distinct from exopod 2. 
Inner spiniform seta of exopod 3 serrate on 



one or both margins, longer and stouter 
than outer, naked spiniform seta. 

Type locality.— Pool #41, 40°37'N, 
121°03'W, Lassen County, Cahfornia. 

Etymology. — The species name is given 
after the State of California, in which the 
type locality is located. 

Habitat description. — The three pools 
(41, 42 and 43 — Long Lake) are relatively 
large for vernal pools (400 X 400 m, 300 
X 300 m and 800 X 200 m respectively at 
time of sampling), shallow (maximum 
depths 0.15-0.6 m), clear, and covered 70- 
90% with aquatic grasses. The electrical 
conductivity was low (30, 60 and 10 
IjlMHO), water temperature 16, 16 and 23 
C, pH 7.7, 7.7 and 8.2, alkalinity 20, 28 and 
8 ppm, and total dissolved solids 10, 30 and 
ppm respectively. 

Discussion and comparisons. — Hespero- 
diaptomus californiensis differs from H. 
schejferi (Wilson, 1953) in the male leg 5: 
the conical process on the coxa of the right 
leg is larger; the inner protrusion on the ba- 
sis of the right leg of H. californiensis is 
rounded, not quadrate as in H. schejferi; 
and the endopods of both left and right legs 
are longer in H. californiensis. In leg 5 of 
the female, H. californiensis has the endo- 
pod extended in a terminal point. The gen- 
ital double somite in H. schefferi has very 
slight lateral projections. 

Hesperodiaptomus victoriaensis is mor- 
phologically closest to H. californiensis. 
The male of H. californiensis differs from 
H. victoriaensis in having the right leg 5 
with longer endopod, and in having the in- 
ner process on the basis larger and placed 
at about midlength rather than at the inner 
proximal corner. In the female of H. victo- 
riaensis, the lateral projections of the gen- 
ital double somite are slightly asymmetri- 
cal, the left projection being directed pos- 
terolaterally rather than dorsally. 

Both H. schefferi and H. victoriaensis 
possess an acute process on the distal mar- 
gin of segment 16 of the right antennule of 
the male. Such a process is lacking in H. 
californiensis. 



VOLUME 109, NUMBER 1 



109 



The differences between H. kiseri and H. 
californiensis are primarily in leg 5 of the 
male: in H. californiensis there is no prox- 
imal protrusion on the inner margin of the 
right basis, and only one protrusion on the 
middle of the inner margin. There is no dis- 
tally placed spiniform projection on the in- 
ner margin of the right exopod 1 . The small 
protrusion on the posterior surface of the 
right exopod 2, present in H. kiseri, is lack- 
ing in H. californiensis. The claw is 
smoothly tapered in H. californiensis, with- 
out angles as in H. kiseri. The lateral pro- 
jections of the genital double somite of the 
female are asymmetrical in H. californien- 
sis, symmetrical in H. kiseri. The thoracic 
wings are not expanded laterally in H. cal- 
iforniensis. 

In H. californiensis, the left lateral pro- 
jection on the female double somite is dor- 
sally directed. The second (middle) uro- 
somite is relatively long. On leg 5 of the 
male, the endopodites are relatively longer 
than in other similar species. 

In the most recent diagnosis of the genus 
Hesperodiaptomus, Borutskii et al. (1992) 
stated that the left antennule of the male is 
like that of the female. However, H. hirsu- 
tus (Wilson, 1953) always, and //. schefferi 
sometimes, display sexual dimorphism in 
setation on some segments (Wilson 1953). 
The observed setation pattern of H. califor- 
niensis was constant, similar in both sexes, 
and the most common pattern found in the 
genus (cf. Wilson 1953). 

Species of the genus Hesperodiaptomus 
are supposed to lack Schmeil's organ (Wil- 
son 1953). Schmeil's organ (Schmeil 1896) 
is a small protuberance of unknown func- 
tion, which in diaptomids if present is lo- 
cated on the posterior surface of leg 2 en- 
dopod 2. If the small transverse ridge in this 
position which was noted in some of the 
males of H. californiensis examined, does 
correspond to Schmeil's organ, this is the 
first reported occurrence of this structure in 
the genus. 

Most species of Hesperodiaptomus occur 
in northwestern North America. A few spe- 



cies are found in eastern Canada and the 
U.S.A., and the ranges of three species ex- 
tend into Siberia (Borutskii et al. 1991). 
The species that are morphologically clos- 
est to H. californiensis occur well north of 
its range. Hesperodiaptomus kiseri is found 
in the State of Washington and in Saskatch- 
ewan (Wilson 1959). Hesperodiaptomus 
schefferi is found in the Pribilof Islands, 
Alaska, and the northern Rocky Mountain 
States (Wilson 1959). Hesperodiaptomus 
victoriaensis has been collected in the 
mountains of southwestern Alberta (Ander- 
son 1967, 1971), near Churchill, Manitoba 
(Hebert 1985; Boileau & Hebert 1988), and 
on Victoria Island, N.W.T. (Reed 1958). 

Two other species of diaptomids, Lepto- 
diaptomus tyrrelli and Hesperodiaptomus 
novemdecimus, occurred in large numbers 
together with H. californiensis in all three 
pools sampled. Many instances of co-oc- 
curring diaptomid species have been re- 
ported and have been much discussed in the 
published literature, including reviews by 
Cole (1961) and Hutchinson (1967). Of 34 
examples of co-occurrence listed by Cole 
(1961), 10 involved congeneric species. 
Hutchinson (1967) observed that co-occur- 
ring diaptomid species tend to be of differ- 
ent sizes and suggested that these size dif- 
ferences may indicate non-overlapping 
feeding niches, and that differences in feed- 
ing would likely be found to be related to 
structural differences. 

Several co-occurrences of species of 
Hesperodiaptomus have been reported. 
Wilson (1953) reported H. wardi (Wilson, 
1953) together with H. novemdecimus. An- 
derson (1971) collected H. shoshone (S. A. 
Forbes, 1893) with H. arcticus, and H. 
shoshone with H. victoriaensis. Hammer & 
Sawchyn (1968) found H. arcticus in the 
same pond as H. kiseri, although the inci- 
dences were separated by a five-day inter- 
val. These authors noted distinct size dif- 
ferences between the two species, but pos- 
tulated mutual exclusion between them. An 
especially interesting example was given by 
Anderson (1967), who found H. shoshone 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



with H. victoriaensis and L. tyrrelli together 
in a pond in Alberta. There, H. shoshone 
individuals averaged 1.6 times longer than 
H. victoriaensis and preyed actively upon 
the much smaller L. tyrrelli. Anderson 
(1967) described the enlarged clawlike se- 
tae of the maxilliped of H. shoshone, which 
were apparently an adaptation for this pred- 
atory activity. The corresponding setae of 
H. victoriaensis were found to be unmodi- 
fied, and this species apparently did not pre- 
date upon the smaller diaptomid. In the case 
of the Lassen County Hesperodiaptomus 
pair, the setae of the mouthparts and max- 
illiped of both species are unmodified. Hes- 
perodiaptomus novemdecimus averaged 
larger than H. californiensis in all three 
ponds, and the two could be reliably sorted 
because of this size difference. In accor- 
dance with Hutchinson's (1967) hypothesis, 
we suggest that differences between these 
species in feeding niches are likely to exist, 
although we have made no direct observa- 
tions. 

Acknowledgments 

This article was prepared during the se- 
nior author's participation in the Mentor- 
ship Program of the Thomas Jefferson High 
School for Science and Technology, Alex- 
andria, Virginia. We thank the collector, Ja- 
mie L. King, University of California, Da- 
vis for providing habitat information and a 
draft of the article by King et al. Jamie King 
and two anonymous reviewers made valu- 
able suggestions on an earlier draft. 

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American Midland Naturalist 32:1-90. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1):112-117. 1996 

Orecturus amplus, a new species 

(Copepoda: Siphonostomatoida: Asterocheridae) 

from an alcyonacean in New Caledonia 

Arthur G. Humes 

Boston University Marine Program, Marine Biological Laboratory, 
Woods Hole, Massachusetts 02543, U.S.A. 

Abstract. — A new species of siphonostomatoid copepod, Orecturus amplus, 
is described from New Caledonia, where it is associated with the alcyonacean 
Siphonogorgia variabilis Hickson. The new species may be distinguished from 
its six congeners by its large size and by detailed features of its external anatomy. 



Copepods are frequent associates of Al- 
cyonacea (Humes 1990). Species of the or- 
der Poecilostomatoida are by far the most 
common copepod associates (nearly 180 
species) of these cnidarians. Those of other 
orders are much less frequently found with 
soft corals (Cyclopoida, 4 species; Harpac- 
ticoida, 1 species; and Siphonostomatoida, 
3 species). Three poecilostomatoids have 
been recorded from the alconacean Siphon- 
ogorgia. In Madagascar, Acanthomolgus 
hians (Humes & Ho 1968) lives with Si- 
phonogorgia pichoni Verseveldt (host orig- 
inally reported as S. pendula Studer) and 
Acanthomolgus longispinifer (Humes & Ho 
1968) with S. pichoni (see Humes & Stock 
1973). In the Moluccas (at Pulau Gomumu, 
south of Obi), Acanthomolgus brevifurca 
Humes, 1990, is associated with S. varia- 
bilis Hickson. 

The relatively small number of siphon- 
ostomatoid copepods reported to date from 
soft corals probably reflects the lack of at- 
tention so far given to these associations. In 
this paper, a siphonostomatoid belonging to 
the genus Orecturus, living with Siphono- 
gorgia in New Caledonia, is described. 

Materials and Methods 

The host alcyonacean colony was isolat- 
ed immediately after collection in a plastic 
bag containing sea water Sufficient 95% 



ethanol was later added to make approxi- 
mately a 5% solution. After 2-3 hours, the 
soft coral was gently rinsed, and the wash 
water poured through a fine net (approxi- 
mately 120 holes per 2.5 cm, each hole ap- 
proximately 225 |xm square). The copepods 
were then recovered from the sediment re- 
tained. 

The copepods were measured and studied 
in lactic acid. The length of the body does 
not include the setae on the caudal rami. 
Dissections were prepared using the wood- 
en slide method described by Humes & 
Gooding (1964). All drawings were made 
with the aid of a camera lucida. 

Order Siphonostomatoida Thorell, 1859 

Family Asterocheridae Giesbrecht, 1899 

Genus Orecturus Humes, 1992 

Orecturus amplus, new species 

Figs, la-g, 2a-i, 3a-i 

Type material. — 7 ? $ , 9 S S from the 
alcyonacean Siphonogorgia variabilis 
Hickson, in 30 m, outside Recif Mtere, near 
Noumea, New Caledonia, 22°20'40"S, 
166°13'55"E, 23 Jul 1971. Holotype ? 
(USNM 274207), allotype 6 (USNM 
274208), and 11 paratypes (4 $9,7 S S) 
(USNM 274209) deposited in the National 
Museum of Natural History, Smithsonian 
Institution, Washington. Remaining para- 
types in the collection of the author. 



VOLUME 109, NUMBER 1 



113 




Fig. 1. Orecturus amplus, new species. Female, a, body, dorsal (scale A); b, body, lateral (A); c, urosome, 
dorsal (B); d, genital area, dorsal (D); e, anal somite and caudal ramus, dorsal (C); f, egg, ventral (B); g, rostral 
area, ventral (B). A, = antennule, Aj = antenna, S = siphon. 



Female. — Body (Fig. la, b) with moder- 
ately broad prosome. Length 1.54 mm 
(1.50-1.56 mm) and greatest width 0.80 mm 
(0.78-0.88 mm), based on 7 specimens in 
lactic acid. Greatest dorsoventral thickness 
0.55 mm. Somite bearing leg 1 fused with 
cephalosome. Somite bearing leg 3 with pos- 
teriorly extended, narrowly rounded epi- 



mera. Somite bearing leg 4 narrow, with 
rounded epimera, overlapped dorsally by ter- 
gum of preceding somite. Ratio of length to 
width of prosome 1.38:1. Ratio of length of 
prosome to that of urosome 2.05:1. 

Somite bearing leg 5 (Fig. Ic) 120 X 264 
|jLm, indented laterally, with scalelike spines 
laterally. Genital double-somite broader 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Orecturus amplus, new species. Female, a, antennule, postero-outer (scale C); b, antenna, outer (C); 
c, siphon, ventral (B); d, mandible, anterior (C); e, maxillule, anterior (C); f, maxilla, posterior (C); g, maxilliped, 
posterior (C); h, leg 1 and intercoxal plate, anterior (E); i, leg 2 and intercoxal plate, anterior (E). 



than long, 117 (xm long in midline, 160 jxm 
long including posterolateral spiniform pro- 
cesses, 231 |jim wide; ratio 0.51:1, taking 
length at midline. Genital areas located dor- 
solaterally near indentation. Each genital 
area with 2 small setae (Fig. Id). Two post- 
genital somites from anterior to posterior 55 



X 172 \xm (length including posterolateral 
processes) and 180 X 160 ixm, longer so- 
mite with small scalelike spines (continuing 
on dorsal and ventral surfaces, but not 
shown in Fig. Ic, e). 

Caudal ramus (Fig. le) 60 X 75 fxm, 
wider than long, ratio 0.8:1. Dorsal seta 70 



VOLUME 109, NUMBER 1 



115 




Fig. 3. Orecturus amplus, new species. Female, a, leg 3 and intercoxal plate, posterior (scale E); b, leg 4 
and intercoxal plate, anterior (E); c, leg 5, ventral (C). Male, d, body, dorsal (A); e, somites of legs 3 and 4, 
dorsal (B); f, urosome, dorsal (E); g, antennule, dorsal (C); h, leg 5, ventral (D); genital somite showing leg 6, 
ventral (D). 



|jLm, with elongate pedicel. Outer lateral 
seta 250 |xm, outermost terminal seta 275 
|xm, innermost terminal seta 340 |xm, and 2 
median terminal setae 460 |xm (outer) and 
625 |xm (inner) swollen proximally. All se- 
tae with long lateral setules. Outer margin 
of ramus with few scalelike spines and pos- 



tero-inner comer with group of slender se- 
tules. 

Dorsal surface of body without visible 
sensilla. 

Entire egg sac not seen, but isolated egg 
oval (Fig. If), 226 X 179 |xm. 

Rostrum (Fig. Ig) weak. Antennule (Fig. 



116 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



2a) 585 |jLm long, 17-segmented. Lengths of 
its segments (measured along their posterior 
nonsetiferous margins): 39 (78 |xm along its 
anterior margin), 26, 83, 26, 25, 4, 21, 34, 
36, 36, 38, 38, 39, 44, 26, 23, and 13 |xm, 
respectively. Formula for armature: 2, 2, 10, 
2, 6, 1, 1, 2, 2, 2, 2, 2, 2, 2 + 1 aesthetasc, 
2, 3, and 5 + 1 aesthetasc. Enlarged seta 
on third segment 74 |jLm long. Certain setae 
on segments 1^ with lateral setules. An- 
tenna (Fig. 2b) with elongate, 1 -segmented 
exopod 57 X 10 |jLm, bearing short inner 
seta and long terminal barbed seta 95 |xm; 
both sides of exopod with setules. Endopod 
with first segment unarmed, second seg- 
ment with 3 short setae and long terminal 
seta 120 ixm long. 

Siphon (Fig. 2c) 935 (xm long, reaching 
almost to ventral edge of intercoxal plate of 
leg 4. 

Mandible (Fig. 2d), maxillule (Fig. 2e), 
and maxilla (Fig. 2f) resembling those of 
congeners (see Humes, 1992, 1993, 1994). 
Maxilliped (Fig. 2g) with basis lacking in- 
ner seta but having small spinules along 
outer margin. Endopod 3-segmented, armed 
with 2,2,1; terminal claw 120 ixm. 

Legs 1-4 (Figs. 2h, i, 3a, b) segmented 
and armed as follows (Roman numerals in- 
dicating spines, Arabic numerals represent- 
ing setae): 

P, coxa 0-1 basis l-I exp I-l; I-l; 11,1,5 

enp 0-1; 0-2; 1,5 
Pjcoxa 0-1 basis 1-0 exp I-l; I-l; 111,1,4 

enp 0-1; 0-2; 1,1,1,3 
Pjcoxa 0-1 basis 1-0 exp I-l; I-l; 11,11,3 

enp 0-1; 0-2; 1,1,3 
P4Coxa 0-1 basis 1-0 exp M; LI; 11,11,3 

enp 0-1; 0-2; 1,1,2 

Leg 1 with inner barbed spine on basis 
52 |xm. 

Leg 5 (Fig. 3c) placed ventrally (as 
shown in Fig. Ic). Suboval free segment 
107 X 70 |jLm, ratio 1.53:1. Five setae from 
outer to inner 75, 65, 36, 25, and 27 |xm. 
Two innermost setae stout, almost spine- 
like, and lacking setules. Other setae with 
lateral setules. Few scalelike spines along 



outer edge of free segment. Adjacent dorsal 
seta, here located ventrally, 52 |xm with lat- 
eral setules. 

Leg 6 represented by 2 small setae on 
genital area (Fig. Id). 

Color of living specimens red, eye red. 

Male.— Body (Fig. 3d) with prosome 
more slender than in female. Length 0.98 
mm (0.94-1.02 mm) and greatest width 
0.43 mm (0.40-0.45 mm), based on 9 spec- 
imens in lactic acid. Greatest dorsoventral 
thickness 0.33 mm. Somite bearing leg 4 
with sharply pointed epimera (Fig. 3c). Ra- 
tio of length to width of prosome 1.69:1. 
Ratio of length of prosome to that of uro- 
some 2.46:1. 

Somite bearing leg 5 (Fig. 3f) 73 X 127 
|xm including processes (length at midline 
62 |jLm). Genital somite 60 X 86 |xm in- 
cluding leg 6 (length at midline 65 |xm). 
Three postgenital somites from anterior to 
posterior 42 X 112 ixm (length at midline 
26 |JLm), 26 X 104 |jLm (length at midline 
16 |xm), and 104 X 112 |xm. 

Caudal ramus 39 X 52 |xm, ratio 0.75:1, 
resembling that of female. 

Dosal surface of body without visible 
sensilla. Urosome with scalelike spines as 
in female. 

Rostrum as in female. Antennule (Fig. 
3g) 1 3-segmented, with segments 8 and 9 
clearly separated (in other species these 
segments tending to be fused). Lengths of 
segments (measured along their posterior 
nonsetiferous margins): 21 (52 jjim along 
anterior margin), 22, 42, 23, 13, 9, 14, 26, 
48, 29, 57, 52, and 48 ixm, respectively. 
Formula for armature: 2, 2, 10 + 3 aesthet- 
ascs, 2 + 1 aesthetasc, 6, 1, 1, 2, 4, 2, 1, 1 
+ 1 aesthetasc, and 5 + 1 aesthetasc. 
Aesthetascs on segments 3, 4, and 10 (these 
present in all males examined) very hya- 
line. Antenna as in female. 

Siphon, mouthparts, and legs 1^ like 
those of female. 

Leg 5 (Fig. 3h) with free segment 50 X 
32 |xm, ratio 1.56:1, otherwise placed ven- 
trally and similar to that of female. 



VOLUME 109, NUMBER 1 



117 



Leg 6 (Fig. 3i) posteroventral flap on 
genital somite bearing 2 slender setae. 

Color of living specimens as in female. 

Etymology. — The specific epithet am- 
plus, Latin meaning large, alludes to the rel- 
atively large size of this species compared 
to congeners. 

Remarks. — Orecturus amplus may be 
distinguished by its size. None of its six 
congeners exceeds a length (in the female) 
of L25 mm, with their average length of 
only 1.11 mm. Selected features for further 
differentiation are as follows. In O. braca- 
tus (Stock & Kleeton 1963) the genital dou- 
ble-somite is rectangular. In O. excavatus 
Humes, 1989, the outer side of segment 1 
of the exopod of leg 1 is excavated and the 
posterior part of the claw of the maxilliped 
is swollen. In O. finitimus Humes, 1993, the 
enlarged seta on segment 3 of the antenna 
is longer than the segment and the basis of 
the claw of the male maxilliped has a low 
prominence. In O. forticulus Humes, 1993, 
the prosome is broad, the maxilla has an 
unusually stout claw, the somite bearing leg 
3 has sharply pointed epimera, and the first 
segment of the exopod of leg 1 has an outer 
thomlike process. In O. grandisetiger Hu- 
mes, 1992, the prosome is broad, the en- 
larged seta on segment 3 of the antennule 
is longer than the segment, the somite bear- 
ing leg 3 has sharply pointed epimera, and 
the basis of the male maxilliped has a weak 
inner protuberance and lacks spinules. In O. 
sakalavicus Humes, 1994, the antennule 
lacks an enlarged seta, the somite bearing 
leg 3 has sharply pointed epimera, and the 
basis of the male maxilliped has an inner 
lobe and lacks spinules. The outer spinules 
on the basis of the maxilliped seen in the 
new species are found only in O. forticulus. 

The color of living colonies of the soft 



coral host, Siphonogorgia variabilis, was as 
follows: slender creamy white stems, small- 
er branches and twigs pale yellow, polyps 
lavender blue to red. The red color of Or- 
ecturus amplus more closely resembles that 
of the polyps than that of other parts of the 
colony, suggesting that the copepods may 
live on or in the polyps. 

Literature Cited 

Humes, A. G. 1989. Acontiophorus excavatus, a new 
species (Copepoda: Siphonostomatoida) asso- 
ciated with the soft coral Dendronephthya (Al- 
cyonacea) in the Indo-Pacific. — Proceedings of 
the Biological Society of Washington 102:916- 
923. 

. 1990. Synopsis of lichomolgid copepods 

(Poecilostomatoida) associated with soft corals 
(Alcyonacea) in the tropical Indo-Pacific. — 
Zoologische Verhandelingen 266:1-201. 

. 1992. Copepoda associated with the thorny 

coral Antipathes (Antipatharia) in the Indo-Pa- 
cific. — Journal of Natural History 26:709-744. 

. 1993. Copepoda associated with gorgona- 

ceans (Cnidaria) in the Indo-Pacific. — Bulletin 
of Marine Science 53:1078-1098. 

. 1994. Copepoda associated with octocorals 

in northwestern Madagascar, including Orectu- 
rus sakalavicus n. sp. from the telestacean Coe- 
logorgia palmosa. — Transactions of the Amer- 
ican Microscopical Society 113:117-126. 

, & R. U. Gooding. 1964. A method for study- 
ing the external anatomy of copepods. — Crus- 
taceana 6:238-240. 

, & J.-S. Ho. 1968. Cyclopoid copepods of the 

genus Lichomolgus associated with octocorals 
of the families Xeniidae, Nidaliidae, and Teles- 
tidae in Madagascar. — Proceedings of the Bio- 
logical Society of Washington 81:693-750. 

, & J. H. Stocii. 1973. A revision of the family 



Lichomolgidae, cyclopoid copepods mainly as- 
sociated with marine invertebrates. — Smithson- 
ian Contributions to Zoology 127:1-368. 
Stock, J. H., & G. Kleeton. 1963. Copepodes associes 
aux invertebres des cotes du Roussillon 3. — 
Acontiophorus bracatus n. sp. un cyclopoide si- 
phonostome associe aux six octocoralliaires. — 
Vie et Milieu 14:551-560. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1): 118-137. 1996 

Records of Enchytraeidae (Annelida: Oligochaeta) from west Florida. 

1. MesenchytraeuSy Cognettia, Bryodrilus, Hemienchytraeus, 

Henlea and Buchholzia 

Brenda Healy 

Department of Zoology, University College, Belfield, 
Dublin 4, Republic of Ireland 

Abstract. — Records of eleven species of Enchytraeidae belonging to six gen- 
era are given for terrestrial habitats in Florida west of Talahassee. All are new 
records for the southern part of North America. A revised diagnosis of Mes- 
enchytraeus is provided and a new species, Mesenchytraeus hamiltoni, is de- 
scribed which is characterised by the occurrence of secondary pharyngeal 
glands in V and a very short spermathecal ectal duct. Four species of Cogettia 
occurred, including two new species: Cognettia floridae which belongs to the 
group of species with three pairs of primary and two pairs of secondary pha- 
ryngeal glands and male organs in the normal position, and a species for which 
the description is incomplete. Bryodrilus novaescotiae, Hemienchytraeus ste- 
phensoni and H. bifurcatus are fully described and the occurrence of the wide- 
spread species Henlea perpusilla, H. ventriculosa and Buchholzia fallax is re- 
corded. 



The species recorded and described in 
this paper occurred in four series of collec- 
tions of Enchytraeidae made in West Flor- 
ida (west of Talahassee) during March 
1984, April-May 1987, March 1988 and 
December 1993. Collection procedures and 
the range of habitats sampled have been de- 
scribed in a previous paper that also con- 
tains comments on faunal composition and 
ecology (Healy 1989). The enchytraeid fau- 
na was found to be rich but patchily dis- 
tributed. Among the 70-75 taxa recognised 
were at least 24 species believed to be new 
to science, a further 10 species new to 
North America and 18 already known from 
the continent but new to Florida. The large 
number of new species and new regional 
records is explained by the fact that there 
have been few studies of enchytraeids in 
North America and almost none in the 
Southeast. The only enchytraeids previous- 
ly recorded from Florida are two species of 
Grania from the marine sublittoral on the 
Atlantic coast (Kennedy 1966, Coates & 



Erseus 1985), Stephensoniella marina from 
the marine littoral on Virginia Key (Coates 
1983) and the freshwater species Barbidri- 
lus paucisetosus from the Appalachicola 
River (Loden & Locy 1980). There are no 
records of terrestrial species. 

This paper records details of 1 1 terrestrial 
species belonging to six of the 13 genera 
found in West Florida. Most of the previ- 
ously known species were originally de- 
scribed from European populations, thus 
some morphological differences are to be 
expected in North American specimens. 
When Florida specimens depart from the 
original descriptions or those of European 
material as given by Nielsen & Christensen 
(1959), these differences are noted. A full 
description of Bryodrilus novaescotiae is 
given because the original description was 
based on a small number of fixed specimens 
only. Hemienchytraeus stephensoni and H. 
bifurcatus are also fully described, extend- 
ing the known range of variability in these 
widespread species. 



VOLUME 109, NUMBER 1 



119 



Materials and Methods 

Samples were collected from a wide 
range of habitats, particularly moist or wet 
humus in woodlands or swamps, and soil 
from capillary zones at the edge of water 
bodies such as bayheads, ponds and 
streams. Enchytraeids were absent or in 
poor condition in dry soils and rare in sub- 
merged substrates where they were gener- 
ally replaced by Tubificidae. The species 
described in this paper occurred at nine lo- 
calities. 

Worms were extracted from soil samples 
using the wet funnel method which uses 
light and heat from light bulbs to expel 
worms from samples into water-filled fun- 
nels (O'Connor 1955, Healy & Rota 1992). 
Some specimens of all species were exam- 
ined live and measurements of live worms 
or their organs were made on worms lightly 
compressed under a cover slip. Other ma- 
terial was fixed in 70% ethanol or Bouin's 
fixative and stored in 70% ethanol. Most 
specimens were stained in paracarmine and 
whole-mounted in Canada Balsam but a 
few unstained mounts were also prepared. 

Type specimens and other material have 
been deposited in the United States Nation- 
al Museum of Natural History, Washington 
D.C. (USNM) and a few specimens are in 
the Royal Ontario Museum (Invertebrate 
Zoology), Toronto (ROMIZ). 

Genus Mesenchytraeus Eisen, 1878 

Mesenchytraeus Eisen, 1878:67. 
Analycus Levinsen, 1884:230. 
Mesenchytraeus Eisen 1879:10, 1904:14- 

17; Michaelsen 1887:369-372, 1889:15- 

16; Cejka 1914:5; Welch 1920:42-44; 

Cernosvitov 1937a:278; Nielsen &. Chris- 

tensen 1959:30-31. 

Type species. — Mesenchytraeus primae- 
vus Eisen, 1878. 

Following the listing of Eisen's species 
M. primaevus, M. mirabilis and M. falci- 
forrnis (Eisen 1878) as species dubia by 
Nielsen & Christensen (1959), Brinkhurst 



& Jamieson (1971) designated Analycus 
Levinsen, 1884 as the correct name for the 
genus on grounds of priority. However, Pip- 
er et al. (1982) re-examined Eisen's mate- 
rial and concluded that the specimens were 
undoubtedly members of Mesenchytraeus 
although their poor condition precluded the 
distinction of specific characters. As the 
original descriptions were deemed ade- 
quate, they therefore restored the name 
Mesenchytraeus designating M. primaevus 
as type species. 

Diagnosis. — Medium to large worms, 
usually white but sometimes yellow due to 
colored coelomocytes or black when the 
epidermis is pigmented. Setae sigmoid, dis- 
tinctly nodulated or with slight swellings. 
Head pore at or near the apex of the pro- 
stomium. Clitellum well developed, some- 
times extends over several segments. Paired 
pharyngeal glands on septa at 4/5 and 5/6, 
secondary pharyngeal glands, when pres- 
ent, ventral in two to five segments. Tran- 
sition between esophagus and intestine 
gradual. Peptonephridia absent. Esophagus 
and intestine without diverticula or append- 
ages. Dorsal vessel originates in, or more 
usually behind, clitellum. Coelomocytes of 
one type, nucleate, rather small for the size 
of the worm, round, oval or spindle-shaped. 
Nephridia with poorly developed interstitial 
tissue, allows the coils of the canal to be 
seen clearly, preseptale consisting of a ne- 
phrostome on a short or long neck, post- 
septale with two or three lobes. Brain rather 
short, often broader than long, its anterior 
border indented or cleft. Paired seminal 
vesicles usually present, sometimes extend- 
ing within the egg sac into post-clitellar 
segments. Sperm funnel small, barrel- 
shaped or cylindrical. Sperm duct often en- 
larged to form an atrium just before it en- 
ters penial bulb. Atrial glands (prostates) 
and penial glands present or absent. Egg sac 
present, often extends through several seg- 
ments behind clitellum. Spermathecae sim- 
ple or with diverticula, free or fused to 
esophagus. 

Remarks. — The above diagnosis is a 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



modified version of those given by Eisen 
(1878, 1879, 1904), and Nielsen & Chris- 
tensen (1959), and takes into account more 
recent descriptions. Nodulated setae were 
considered diagnostic by Nielsen & Chris- 
tensen but were not mentioned by Eisen, 
either in his generic diagnoses or in any of 
his detailed species descriptions. Subse- 
quent reviews of the genus (e.g., Michael- 
sen 1887, 1889; Cejka 1914, Welch 1920, 
Cemosvitov 1937a) similarly omitted men- 
tion of the character. Re-examination of the 
material described in Eisen's 1878 paper by 
Piper et al. (1982) confirmed the presence 
of noduli in all three species and it must be 
assumed that the character was overlooked 
by early workers. The nodulus may be very 
weak, however, and may be represented by 
little more than a slight swelling, as in the 
new species described below, making it dif- 
ficult to recognise for an inexperienced 
worker. A nodulus is unusual in enchy- 
traeids and is present in only one other ge- 
nus. However, reliance should not be placed 
on this character when constructing or us- 
ing keys to genera. Other modifications of 
existing diagnoses concern the postseptal 
regions of the nephridia, which Nielsen & 
Christensen describe as bilobed but which 
frequently have three lobes, and the egg 
sac, which Eisen describes as single and 
median but which may extend backward as 
two distinct sacs. Eisen (1904) described 
the penial bulb as a muscular cushion con- 
taining radiating muscular strands, which in 
some species are powerful and usually con- 
tain internal glands (penial glands), but the 
bulb structure is variable and the whole 
structure may be quite compact. Although 
not always present, an atrium, formed by 
expansion of the sperm duct at the entrance 
to the penial bulb, sometimes with attached 
glands, is unique in the Enchytraeidae. 
None of the characters proposed by Eisen 
(1878, 1879) as grounds for erecting the 
new genus has proved to be useful; these 
are: the presence of sperm balls or sper- 
matophores in the coelom (not recorded in 
any other enchytraeid genus but only pres- 



ent in some Mesenchytraeus species), brain 
deeply divided in front (not unique to Mes- 
enchytraeus, indentation sometimes shal- 
low), and an unusually short vas deferens 
(usually short but not always exceptionally 
so). Cardiac glands (Herzkorper of Mi- 
chaelsen), mentioned by early workers, are 
internal structures in the dorsal vessel, not 
easily detected in whole mounts. In prac- 
tice, the most reliable character that distin- 
guishes Mesenchytraeus is the nephridium 
with its free nephrostome (also found in 
some other genera), poorly developed in- 
terstitial tissue and lobed postseptale. In 
live worms, the nephridia have a very dis- 
tinctive appearance that makes Mesenchy- 
traeus specimens instantly recognisable. 

Distribution. — Eisen (1904) described 18 
species of Mesenchytraeus from western 
parts of North America (mainly Alaska and 
California) and a further ten species have 
been described from western regions by 
other workers; in contrast, only three spe- 
cies are recorded from the eastern part of 
the continent (Tynen 1975). The genus also 
appears to be well represented in Siberia 
(Nurminen 1973a, Piper et al. 1982, Timm 
& Popchenko 1978, Timm 1994) while 
only eleven species are recognised in Eu- 
rope. Very few species have been found in 
the tropics. The genus thus appears to have 
its present centre of distribution in northern 
regions of the western Nearctic and eastern 
Palearctic. The poor representation in West 
Florida, where only one species was at all 
frequent, supports this view. 

North America, South America, Green- 
land, Europe, Spitzbergen, Siberia, Japan, 
Antarctic, ?Africa. 

Mesenchytraeus hamiltoni, 

new species 

Fig. 1 

Material examined. — Holotype: USNM 
170722 whole mounted specimen from 
edge of "sewage pond". University of West 
Florida, Pensacola, collected by author, De- 
cember 1993. Paratypes from the type lo- 



VOLUME 109, NUMBER 1 



121 




Fig. 1. Mesenchytraeus hamiltoni, new species. A, anterior segments, dorsal view; B, dorsal setal bundle; 
C, brain; D, spermatheca of live worm, contracted; E, spermatheca of live worm, expanded; F, coelomocytes; 
G, dorsal clitellar gland cells and clear interspaces; H, sperm funnel, live specimen; I, sperm funnel, fixed 
specimen; J, male pore and associated structures; K, nephridium. am ampulla, at atrium, br brain, dv dorsal 
vessel, ed ectal duct, hp head pore, mp male pore, pb penial bulb, phb pharyngeal bulb, pph primary pharyngeal 
gland, pr prostate, sd sperm duct, sp spermatheca, sph secondary pharyngeal gland. 



cality: USNM 170723-170726, two stained 
and two unstained whole mounts, one with 
clitellum partly removed. Other material: 
seven whole mounts in author's collection. 
Approximately 16 live mature specimens 
examined. 

Type locality. — Wet or saturated peaty 
sand with moss and plant roots at edge of 
a pond receiving clean water from a sewage 
treatment plant, water level constant. pH 
4.5-5.3. 

Etymology. — The species is dedicated to 
Professor Paul Hamilton of the University 
of West Florida who introduced me to many 
interesting and rewarding localities. 

Description. — Medium to large, white, 
yellowish or pink worms; live specimens 



18-30 mm, 0.7 mm in diameter. Fixed 
specimens often much contracted, mature 
individuals measuring 8-22 mm with a 
prominent clitellum up to 1.2 mm in di- 
ameter. Segments in mature specimens 55- 
86 (X = 66, SD 7.9, n = 11), 55-75 in 
submature ones. Setae sigmoid with nodu- 
lus poorly developed or absent, without en- 
tal hook (Fig. IB), 108-120 [xm in the pre- 
clitellar region, 120-144 ixm in posterior 
segments. Lateral bundles with three or four 
setae, ventral bundles with 3-12 in precli- 
tellar segments, 3-1 1 behind the clitellum, 
setae about equal in length within bundle. 
Setae absent dorsally in XII and XIII, oc- 
casionally present ventrally in XII, reduced 
in number or absent ventrally in XIII. Cu- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



taneous glands in two or three rows at about 
the level of the setae, or scarcely visible. 
Clitellum extends from V2XI to % or all of 
XIII with numerous small gland cells con- 
taining coarse granules, irregularly scat- 
tered with clear interspaces, that mask in- 
ternal organs in live as well as stained spec- 
imens (Fig. IG). Gland cells may be almost 
continuous ventrally in fully mature indi- 
viduals. Head pore at, or near, tip of pro- 
stomium. 

Primary pharyngeal glands on 4/5 and 
5/6 free dorsally, either unlobed or with two 
or three lobes; secondary glands, each with 
several lobes, situated ventrally in V, VI, 
VII and usually in VIII (Fig. lA). A few 
specimens had either one or two glands in 
IX and one individual possessed a lobed 
gland unilaterally in X (Fig. lA). Two small 
bulbs on dorsal posterior border of the phar- 
ynx in IV (Fig. lA). Chloragocytes small, 
brown, present from IV, form a dense layer 
from VIII, 12-16 cells across the intestine 
in the preclitellar region in mounted speci- 
mens. Esophagus merges with intestine 
without increase in diameter, intestine en- 
larges behind origin of dorsal vessel. Coe- 
lomocytes 14-30 ixm, oval or spindle- 
shaped, with rather coarse granules and a 
weakly staining nucleus (Fig. IF). Dorsal 
vessel originates in XI-XIII with dilatations 
in VII-XII and conspicuous branches in II 
and III (Fig. lA); bifurcation anterior to 
brain. Blood faintly or distinctly red. Brain 
about as broad or slightly broader than 
long, about 80 fxm in length, truncated or 
slightly indented posteriorly, deeply indent- 
ed on its anterior border (Fig. IC). Nephrid- 
ial nephrostome elongated, on a long neck, 
postseptale with two or three lobes; efferent 
duct arises mid-ventrally (Fig. IK). First 
nephridia at 6/7, occasionally at 5/6. 

Testes large and spongy in mature 
worms, appear brown by transmitted light 
in living specimens. Seminal vesicles 
paired, confined to XI or extend into X. 
Sperm funnels small, barrel- shaped, 120- 
160 |jLm in length, about 1.5 times as long 
as wide, with a raised collar about as wide 



as the funnel in fixed specimens, appearing 
slightly narrower in live worms (Fig. IH, 
I). Sperm duct 14-16 |xm wide, about ten 
times length of funnel, loosely coiled in 
XII. Male openings irregular or semicircu- 
lar slits. Each sperm duct widens distally to 
form a thick-walled, fusiform atrium that 
opens centrally into penial bulb; three large, 
slightly lobed glands attached to bulb (Fig. 
IJ). Two egg sacs develop asymmetrically 
on 11/12 as pouches, each containing a 
string of oocytes and eggs, reaching to 
XVIII on one side. Ovarian tissue diffuse, 
extending back in egg sacs which develop 
before sperm funnels and ducts. Up to five 
mature eggs present at a time. Spermathe- 
cae simple, without diverticula, each con- 
sists of an elongated ampulla that tapers en- 
tally to merge with esophagus in V and ec- 
tally to a short ectal duct, only a little longer 
than wide, scarcely longer than thickness of 
body wall (Fig. ID, E). Ampullae expand 
and contract and occasionally pulsate. 
When contracted, wall becomes rugose or 
folded internally (Fig. ID). Only small 
amounts of sperm, arranged lengthwise, in 
ampulla. Spermathecae develop early and 
are fully formed in specimens only half the 
final size and in which there are no signs 
of male organs or oocytes. 

Remarks. — Among the ten or so de- 
scribed species of Mesenchytraeus with 
spermathecae attached to the esophagus in 
V, but which lack diverticula, M. hamiltoni 
is unique in the shortness of the ectal duct 
of its spermatheca and in the presence of 
secondary pharyngeal glands in V. These 
features are deemed sufficient to character- 
ise the species although details of structures 
associated with the male opening, often an 
important diagnostic character in the genus, 
have not been described for most of the 
species with which it might be compared. 
Mesenchytraeus hamiltoni most resembles 
M. glandulosus (Levinsen, 1884) which has 
five pairs of secondary pharyngeal glands 
but has a spermathecal ectal duct equal in 
length to one third of the ampulla, lacks 
secondary pharyngeal glands in V and has 



VOLUME 109, NUMBER 1 



123 



a nephridial funnel without a neck. The 
number of pharyngeal glands was variable 
for individuals of the same size and stage 
of maturity. 

Although the species was plentiful at the 
type locality and at several other sites, ma- 
ture specimens were rare. None was found 
in March 1984, only four in April-May 
1987, three in March 1988 and seven in De- 
cember 1993. A low proportion of adults in 
populations is common in the genus (per- 
sonal observation). 

Habitat. — Mesenchytraeus hamiltoni 
was common in the Pensacola region and 
was sometimes the dominant enchytraeid in 
wet substrates. It occurred in a variety of 
habitats including woodland leaf litter in 
valleys and swamps. Sphagnum and pitcher 
plant (Sarracenia) bogs, and saturated soils 
at the edge of water bodies where the high- 
est densities occurred. At the latter sites 
they were often accompanied by Tubifici- 
dae. The pH range was 3.9-5.8. 

Distribution. — In and around the campus 
of the University of West Florida, Pensa- 
cola; Black water Forest, at the edge of a 
swamp; Avalon Peninsula, edge of a pitcher 
plant (Sarracenia) bog; Lakewood, among 
Sphagnum and grassroots at edge of lake. 
Not found in the east of the study area. 

Mesenchytraeus sp. 

Material examined. — Seven live, imma- 
ture specimens. 

Remarks. — A somewhat yellowish spe- 
cies was present at several hardwood forest 
sites but no fully mature specimens were 
available for a definitive identification. 
Specimens had two pairs of primary and 
two pairs of secondary pharyngeal glands 
and spermathecae communicating with the 
esophagus in V, each with a pair of elon- 
gated diverticula. The species resembles M. 
beumeri (Michaelsen, 1886) in general ap- 
pearance. 

Habitat and distribution. — In hardwood 
leaf litter. Occasional at the University of 
West Florida, Pensacola and in Torreya 



State Park; common at Tall Timbers Forest 
Research Station. 

Genus Cognettia 
Nielsen & Christensen, 1959 

Cognettia species are recognised by sig- 
moid setae, without nodulus, in bundles of 
two or three; nephridia with preseptale con- 
sisting of funnel only and an antero- ventral 
or mid-ventral origin of the efferent duct; 
absence of peptonephridia; absence of 
esophageal and intestinal diverticula; dorsal 
vessel originating in or behind clitellum; a 
simple penial bulb and a simple spermathe- 
ca, not attached to the oesophagus. Testes, 
ovaries and male pores may be displaced 
forward by up to four segments. 

Distribution. — The only record of Cog- 
nettia in North America is for C. glandu- 
losa from Montreal, Canada (Nurminen 
1973b). The genus has proved so far to be 
characteristic of wet, acid soils in cold or 
cold- temperate lands and members are usu- 
ally the dominant enchytraeids in tundra 
and coniferous forests of northern Europe. 
However, the discovery of four species in 
Florida shows that Cognettia species can 
survive in a warm climate while the pres- 
ence of two new species suggests that North 
America, like Europe, may have its own en- 
demic species. 

Canada, Greenland, Europe, Spitzbergen, 
Siberia, Japan, Ecuador, Antarctic. 

Cognettia floridae, new species 
Fig. 2 

Material examined. — Holotype: USNM 
170727, stained whole mount, Edward Ball 
Nature Trail, University of West Florida, 
Pensacola, a hardwood, bayhead swamp, 
collected by the author April 1987. Para- 
types from the type locality and other sites 
in the University of West Florida: USNM 
170728-170732, two stained and two un- 
stained whole mounts. Other material: 
ROMIZ 13242-13243, stained whole 
mounts; 1 1 whole mounts in the author's 
collection; 42 live specimens examined. 



124 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

30 nm 
pph dv 





Fig. 2. Cognettia floridae, new species. A, anterior segments, lateral view; B, dorsal clitellar gland cells and 
clear interspaces; C, brain; D, spermatheca of live worm; E, spermatheca of fixed specimens; F, nephridium; G, 
coelomocytes; H, setae; I, sperm funnel, live worm, am ampulla, br brain, dv dorsal blood vessel, ec ectal glands, 
ed ectal duct, mu muscle; pph primary pharyngeal gland, sg secondary pharyngeal gland, sp spermatheca. 



Type locality. — In moist, wet or saturated 
sandy humus under Taxodium and hard- 
wood trees in a partly tidal, freshwater bay- 
head swamp; some areas subject to periodic 
flooding. pH 3.9-4.7. 

Etymology. — Named after the State of 
Florida, genitive case. 

Description. — Live specimens greyish- 
white, 7-9 mm, fixed specimens 5-9 mm, 
diameter 0.5-0.52 mm, 0.56 mm at clitel- 
lum. Segments (29)39-46 {X 40.9, SD = 
3.6, n = 23). Setae without ental hook (Fig. 
2H), 80-96 |xm in preclitellar region, 80- 
104 |xm in posterior segments, three per 
bundle in all regions, occasionally only 
two, setae absent in XII. Cutaneous gland 
cells small, in numerous rows or more or 
less scattered. Clitellum prominent in live 
worms, extending over XII-XIII with 
squarish gland cells arranged in transverse 
rows. In stained mounts, the gland cells, 
which are packed with coarse granules, are 



irregularly scattered with clear interspaces, 
absent mid-ventrally (Fig. 2B). Head pore 
at 0/1. 

Three pairs of primary pharyngeal 
glands, all free dorsally, without ventral 
lobes, and two pairs of compact secondary 
glands, situated ventrally in V and VI (Fig. 
2A). Esophagus merges gradually with in- 
testine from 6/7. Chloragocytes present 
from IV, forming a dense layer from VI, 
10-12 across intestine in preclitellar region 
in compressed specimens. Dorsal vessel 
arises at 13/14, occasionally in XIV or XV, 
anterior vessel bifurcates at level of 0/1. 
Blood colorless in specimens from drier 
habitats, faintly or distinctly red in those 
from wet substrates. Coelomocytes round 
or oval, finely granular with a prominent 
nucleus (Fig. 2G), 14-32 fxm long, about a 
quarter of length of setae. Nephridial ne- 
phrostome on a short neck, long efferent 
duct arises antero- ventrally or mid-ventrally 



VOLUME 109, NUMBER 1 



125 



(Fig. 2F). First nephridium at 6/7. Brain 
160-200 |xm long, 1.5-2 times longer than 
wide, deeply incised posteriorly (Fig. 2C). 

Seminal vesicle unpaired, dorsal, con- 
fined to XI. Sperm funnels small, cylindri- 
cal, in live worms about 100 fxm long, four 
or five times longer than wide, equal to 
about V4-V3 diameter of live worm (Fig. 21). 
In fixed specimens, length usually about Va 
of worm diameter. Collar slightly narrower 
than funnel. Sperm duct long and narrow, 
diameter 6 |Jim, coiled in XII, opens through 
a compact penial bulb, greatest diameter 
about 90 |xm, which occupies V^ to V^ of the 
diameter of clitellum in mounted speci- 
mens. One, or usually two, eggs present at 
a time. Spermathecae confined to V. Ectal 
ducts thick-walled, about four times length 
of ampullae, each with a mass of fused, 
elongate cells on anterior face, near opening 
(Fig. 2D, E). Ampulla dorsal to oesophagus 
in V, thin- walled, 35-50 fjim long, 1 .2 times 
longer than wide, with most of sperm ar- 
ranged lengthwise and extending into prox- 
imal part of ectal duct in most specimens 
(Fig. 2D, E). 

Remarks. — Cognettia floridae belongs to 
a group of species that comprises C cog- 
nettii (Issel, 1905), C. lapponica Nurminen, 
1965, C. hibernica Healy, 1975 and C. zic- 
sii Dozsa-Farkas, 1989 all of which repro- 
duce sexually and have reproductive organs 
in the normal position for Enchytraeidae. It 
resembles C lapponica, C. hibernica and 
C. zicsii in having three pairs of primary 
pharyngeal glands and two pairs of second- 
aries and spermathecae confined to V but it 
is unique in the asymmetrical ectal swell- 
ings on the spermathecal ducts. 

Habitat. — The species was common in 
and around the campus of the University of 
West Florida and plentiful at two other lo- 
calities. It was found in a range of sub- 
strates including sandy peat and peaty sand. 
Sphagnum, coniferous and broad-leaved 
leaf litter, finely divided humus and among 
roots of aquatic plants at the edge of water 
bodies. All sites were wet, saturated or 
flooded, with pH levels 3.9-4.5, among the 



lowest for the region. Mature individuals 
were common. 

Distribution. — Pensacola, campus of the 
University of West Florida; Avalon Penin- 
sula, pitcher plant (Sarracenia) bog; Lake- 
wood, edge of a lake. Not found in the east- 
em part of the study area. 

Cognettia sp. 
Fig. 3 

Material examined. — Two live mature 
specimens collected in April 1984 were 
available for the following description but 
both were damaged during examination and 
no further mature individuals could be 
found. The remaining material consists of 
one stained, whole-mounted, submature 
specimen, USNM 170732, three stained 
whole mounts of immature specimens 
USNM 170733-170735 and ten whole 
mounts of immatures in the author's collec- 
tion. Approximately 30 live, immature in- 
dividuals were also examined. 

Description. — Medium sized, rather slen- 
der worms, creamy white, internal organs 
partly masked by abundant coelomocytes in 
live worms. Length of live, mature speci- 
mens 10-12 mm; live, non-fragmenting im- 
matures reach 20 mm. Maximum length of 
fixed immature specimens 12 mm, diameter 
0.28-0.37 mm. Segments 50 and 58 in the 
mature specimens but reach 64 in immatu- 
res. Setae without ental hook, 68-84 \xm in 
all regions, three ventrally, three or some- 
times two laterally. Setae absent from IX or 
X in mature individuals. Clitellum only 
slightly raised, extends over IX-^/iX (over 
X-ViXI in submature specimen), gland cells 
irregularly scattered or partly in transverse 
rows (Fig. 3D). Male pores in X or XI. Cu- 
taneous glands small, in numerous rows or 
scarcely visible. Head pore at, or just an- 
terior to 0/1. 

Five pairs of unlobed, primary pharyn- 
geal glands in large individuals but fifth 
pair often small, developing unilaterally or 
absent (Fig. 3G). Only four pairs in mature 
worms. Anterior glands may be united dor- 



126 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ec 







am 






S5*?« 






10 urn 




200 ^m 



Fig. 3. Cognettia sp. A, spermatheca, live worm; B, coelomocytes; C, sperm funnel, live worm; D, clitellar 
glands; E, preclitellar nephridium; F, postclitellar nephridium; G, anterior segments, lateral view, am ampulla, 
br brain, ed ectal duct, eg ectal gland, mu muscle, ph pharyngeal gland, nc ventral nerve cord. 



sally while posterior ones are usually free. 
Secondary pharyngeal glands absent. 
Esophagus merges gradually with intestine. 
Chloragocytes present in a dense layer from 
behind last pair of pharyngeal glands i.e., 
in VIII or IX, 10-14 across the intestine in 
compressed specimens. Dorsal vessel from 
XIV or XV, anterior bifurcation just behind 
0/1. Blood colorless or faintly red. Coelom- 
ocytes 20-30 |jLm, round or oval with a 
sharp outline, packed with fine granules that 
make them appear brown by transmitted 
light in living worms (Fig. 3B). They are 



usually very abundant, often so densely 
packed as to mask internal organs. Nephrid- 
ia with free nephrostome and elongate pre- 
septale, efferent duct long, arising antero- 
ventrally or mid-ventrally (Fig. 3E, F). 
Brain about 1.5 times longer than broad 
with a truncate or sinuous posterior border. 
Seminal vesicle present, unpaired. Sperm 
funnels cylindrical, about four times longer 
than wide, each with narrower, scarcely 
raised, collar (Fig. 3C). Sperm ducts long 
and narrow, opening at compact penial 
bulbs. Spermathecal ectal ducts long, slen- 



VOLUME 109, NUMBER 1 



127 



der, without swellings, leading to ovoid am- 
pullae in VII (Fig. 3A). One large ectal 
gland anterior to each spermathecal open- 
ing. 

Remarks. — The above description is in- 
complete owing to the shortage of mature 
specimens but the distinctive appearance of 
live individuals made them easily recognis- 
able in samples, even as immatures, and 
there seems little doubt that this is a new 
species. Many worms were regenerating an- 
terior or posterior segments and the species 
obviously reproduces principally by frag- 
mentation; sexual individuals were rare in 
the population. The species is close to C. 
sphagnetorum (Vejdovsky, 1878) which re- 
produces in the same way and which also 
has four or five pairs of primary pharyngeal 
glands and no secondary glands. Cognettia 
sphagnetorum, however, has a chamber 
containing a ring of sperm at the junction 
between the ectal duct of the spermatheca 
and the ampulla and more anterior male 
openings (in VIII or IX). Other Cognettia 
with displaced reproductive organs, namely 
C. glandulosa (Michaelsen, 1888), C. paxi 
(Moszyiiski, 1938) and C. anomala (Cer- 
nosvitov, 1928), all have secondary pharyn- 
geal glands. While there are no important 
diagnostic differences between immature 
forms of C. sphagnetorum and the present 
species, the abundant coelomocytes with 
sharp outline and densely granular cyto- 
plasm give the Florida species a distinctive 
appearance. 

Habitat and distribution. — University of 
West Florida, Pensacola, in the drier parts 
of a bayhead swamp and in an adjoining 
ravine with hardwoods. Common in moist, 
but not wet, leaf litter and humus, pH 3.6- 
4.5, above the zone subject to periodic 
flooding. 

Cognettia 1 sphagnetorum 
(Vejdovsky, 1878) 

Pachydrilus sphagnetorum Veydovsky 1878: 

304. 
Cognettia sphagnetorum Nielsen & Chris- 



tensen 1959:42-43, figs. 28, 29; Kas- 
przak 1986:124-125, figs. 332-334; 
Chalupsky 1992:142, fig. 10. 

Material examined. — Two stained, whole- 
mounted immature specimens in the author's 
collection. Three live immature specimens 
examined. 

Remarks. — The absence of mature indi- 
viduals makes a definitive identification im- 
possible, but specimens resembled C. 
sphagnetorum from Ireland in general ap- 
pearance although they had only four pairs 
of pharyngeal glands. Nielsen & Christen- 
sen (1959) give five pairs for Danish ma- 
terial, except in mature individuals, but I 
have found in Irish populations that the fifth 
pair is often absent in fragmenting individ- 
uals. Fragmentation is the normal method 
of reproduction in C. sphagnetorum and 
sexual individuals are always rare. Material 
from Florida included one specimen with a 
developing clitellum and one which ap- 
peared to be a juvenile, with only 20 seg- 
ments, although according to Christensen 
(1959) the species does not seem to be ca- 
pable of producing viable eggs. 

Habitat and distribution. — Lakewood, 
among grass roots and leaf litter at the edge 
of a lake, pH 4.2^.4. Europe, Iceland, 
Spitzbergen, Antarctic. (Some of these re- 
cords are based on immatures only.) 

Cognettia Iglandulosa 
(Michaelsen, 1888) 

Pachydrilus sphagnetorum var. glandulosus 
Michaelsen, 1888:483. 

Marionina glandulosa Issel 1905:455^56, 
fig. 3. 

Enchytraeoides glandulosus Cernosvitov 
1928:16-17, PI. 1, figs. 10, 11. 

Cognettia glandulosa Nielsen & Christen- 
sen 1959:43-44, fig. 30; Kasprzak 1986: 
125-126, figs. 335-337. 

Material examined. — USNM 170736— 
170737, two stained, whole-mounted im- 
mature specimens; two whole mounts in the 
author's collection; five live, immature in- 
dividuals examined. 



128 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





i^o"^> 



Fig. 4. Bryodrilus novaescotiae. A, anterior segments, dorsal view; B, spermatheca, live worm; C, sper- 
mathecal pore, surface view; D, setal bundle; E, preclitellar nephridium; F, postclitellar nephridium; G, brain; 
H, coelomocytes; I, clitellar glands, dorsal view; J, sperm funnel, live worm; K, male pore, ventral view, am 
ampulla, br brain, ch chloragocytes, dv dorsal blood vessel, ed ectal duct, ec ectal glands, en ental duct, eg 
esophageal gland, sp spermatheca. 



Remarks. — The distinctive feature of this 
species is the presence of five pairs of pri- 
mary and five (occasionally four) pairs of 
secondary pharyngeal glands. Immature 
specimens are commonly identified on the 
basis of this character alone. The sperma- 
theca is also distinctive. No sexual individ- 
uals were found, however, during this sur- 
vey, thus a positive identification is not pos- 
sible. Specimens resembled C. glandulosa 
from Ireland in general appearance. The 
presence of regenerating fragments indicat- 
ed that individuals were reproducing by 
fragmentation. 

Habitat and distribution. — Ponce de 
Leon, leaf litter on floodbank of stream is- 
suing from limestone, pH of litter 5.34. 
Canada, Greenland, Iceland, Europe, Sibe- 
ria, Antarctic. 

Genus Bryodrilus Ude, 1892 

Diagnostic characters of the genus are 
sigmoid or curved setae, without nodulus. 



of different sizes within a bundle; four 
esophageal diverticula in VI; peptonephri- 
dia and intestinal diverticula absent; dorsal 
vessel originates within or behind the cli- 
tellum; coelomocytes of uniform size and 
shape; preseptale of nephridium consists of 
funnel only, efferent duct arises anteriorly, 
near the septum C'Henlea type") or medi- 
ally; compact penial bulb; simple sperma- 
thecae whose ental ducts usually unite be- 
fore communicating with the dorsal wall of 
the esophagus. 

Distribution. — So far confined to the 
northern Holarctic, the majority of species 
being recorded from North America and Si- 
beria. Alaska, Canada, Greenland, Europe, 
Siberia. 

Bryodrilus novaescotiae 

Bell, 1962 

Fig. 4 

Bryodrilus novaescotiae Bell, 1962:169— 
171, PI. iv, figs. 1-9. 



VOLUME 109, NUMBER 1 



129 



Material examined. — Three stained 
whole mounts USNM 170746-170748; 
three stained whole mounts ROMIZ 13241, 
13244-13245, 17 whole mounts in the au- 
thor's collection. Approximately 30 live 
specimens examined. Type material not lo- 
cated. 

Description of new material. — Medium 
to large species, white or yellowish due to 
color of chloragogen tissue and abundant 
coelomocytes. Length variable; live speci- 
mens 10-20 mm, reach 30 mm when 
stretched; fixed specimens 8-16 mm, di- 
ameter 0.7-0.8 mm, slightly wider at clitel- 
lum. Segments (37)45-64 (X = 54.2, SD = 
5.5, n — 19). Setae sigmoid, without ental 
hook, three or four in lateral bundles, five 
or six (occasionally seven) in anterior ven- 
tral bundles, four to six ventrally behind cli- 
tellum (Fig. 4D). Setae diminish in size 
from outside of bundle to mid-line and 
measure 50-100 |xm in preclitellar region, 
112-128 |Jim in posterior segments. Cuta- 
neous gland cells in about ten rows anteri- 
orly, more noticeable at level of setae, eight 
or nine rows per segment behind clitellum. 
Clitellum extends over XII and %-% XIII, 
only slightly raised, with numerous small, 
coarsely granular, irregularly scattered 
gland cells and polygonal interspaces (Fig. 
41). Ventrally, gland cells nearly continu- 
ous. Head pore a transverse slit just anterior 
to 0/1. 

Three pairs of lobed pharyngeal glands, 
all free dorsally and with ventral lobes (Fig. 
4A). Four esophageal glands in VI, closely 
applied to the esophagus, situated dorso-lat- 
erally and ventro-laterally, each with five to 
seven regular lobes (Fig. 4A), apparently 
solid, no internal canals seen. Chloragocy- 
tes small, average 10-15 in number across 
intestine in compressed specimens, form a 
dense layer from 6/7 at point where gut 
widens gradually. Coelomocytes numerous, 
round or oval, finely granular, nucleus not 
visible in live worms but distinct when 
stained, diameter (16)20-42 jxm (Fig. 4H). 
In live individuals, coelomocytes appear 
brown by transmitted light. Dorsal vessel 



originates at 11/12 or sometimes 12/13, 
with segmental dilatations in VI-XI, ante- 
rior bifurcation at 0/1. Blood colorless. Ne- 
phridia with small, free nephrostome and 
elongate preseptale, postseptale with long, 
narrow efferent duct that arises antero-ven- 
trally in preclitellar region, mid- ventrally in 
posterior segments (Fig. 4E, F). First ne- 
phridium at 6/7. Brain usually about 1.5 
times longer than wide but sometimes 
shorter with a sinuous posterior border and 
straight or indented anterior border (Fig. 
4G). 

Testes small, oval and compact with a 
smooth outline. Seminal vesicle unpaired, 
confined to XL Sperm funnels cylindrical 
or taper distally, each with a distinct collar 
of about same width as funnel (Fig. 4J), 
length in live specimens 150-180 |xm, three 
or four times longer than wide, up to five 
times longer when stretched, equal to Va to 
y-i of diameter of worm. In mounted speci- 
mens, funnel has a smooth outline and is 
usually bent into a semicircle. Sperms duct 
long and narrow, coiled in XII, about 6 |xm 
in diameter; each opens through a large but 
compact penial bulb, about 60 [xm in lon- 
gitudinal diameter but sometimes smaller. 
In fixed specimens, bulb occupies about Va 
of diameter of clitellum. Male openings 
semicircular slits (Fig. 4K). One egg pres- 
ent at a time. Spermathecae large, more or 
less confined to V (Fig. 4B). Ectal duct of 
spermatheca composed of two parts roughly 
equal in length, a stout, thick-walled section 
communicating with one side of ampulla, 
and a large, bulbous, asymmetrical mass of 
cells surrounding a narrow chamber that 
opens to the outside through a straight or 
curved slit at 4/5, surrounded by a protrud- 
ing ring of large cells forming a rosette at 
the surface up to 190 ixm across (Fig. 4C). 
Ampulla thin-walled, roughly pear-shaped 
in live specimens but contracting to become 
more spherical when fixed. Ampullae taper 
entally, ental ducts usually unite just before 
communicating with esophagus at 5/6 or in 
anterior part of VI (Fig. 4B), but occasion- 
ally communicate separately. Ampullae and 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ental ducts contain abundant sperm, mostly 
arranged lengthwise. 

Remarks. — The distinctive character of 
B. novaescotiae is its spermathecae. The 
large swellings near the openings of the ec- 
tal ducts are not present in any other mem- 
ber of the genus. Bell (1962) also consid- 
ered the solid esophageal glands to be 
unique as they are hollow in other species. 
His specimens differed chiefly in the form 
and dimensions of the spermathecae, each 
of which had a large, hemispherical cham- 
ber in the ectal bulb, a smaller "spherical" 
ampulla and a longer common ental duct. 
He mentioned two ectal glands in addition 
to the mass of fused cells around the orifice, 
but these are not shown in his figure (Bell 
1962, fig. 3). Bell also noted a hemispher- 
ical chamber within the penial bulb which 
could not be detected in mounted speci- 
mens from Florida. Other differences in 
Bell's specimens are a shorter brain, about 
half as long as wide, irregular- shaped coe- 
lomocytes (possibly resulting from poor 
fixation) and nephridia with a large presep- 
tale and an efferent duct that arises termi- 
nally but is folded back to give the impres- 
sion of leaving subterminally or even near 
the septum. This latter character cannot be 
considered significant since in sectioned 
material it can be difficult to tell whether 
the distal portion of the nephridial duct is 
folded inside or outside the sheath of the 
postseptale. Bell did not mention lobes on 
the esophageal glands, which are not pres- 
ent in other known species of Bryodrilus, 
the dimensions of the sperm funnel, or the 
presence of a seminal vesicle. 

Habitat and distribution. — University of 
West Florida, Pensacola, in drier parts of a 
hardwood swamp and an adjoining wooded 
ravine, mainly above the level of periodic 
flooding. Soil a moist, spongy humus with 
numerous fine roots, pH 3.5-4.5. Canada 
(Nova Scotia). 

Genus Hemienchytraeus 
Cernosvitov, 1934 

Hemienchytraeus species possess a me- 
dian, unpaired, bifurcated peptonephridium 



arising from the dorsal wall of the pharynx, 
which is unique in the Enchytraeidae. Other 
useful characters are setae in pairs; absence 
of esophageal or intestinal diverticula; ne- 
phridia with large preseptale enclosing the 
nephrostome; long sperm ducts usually 
coiled into a spirals; small, compact penial 
bulbs and free spermathecae each of which 
has an ectal duct without glands at the 
opening and a simple ampulla without di- 
verticula. 

Distribution. — Hemienchytraeus species 
were the most abundant and widespread en- 
chytraeids in West Florida, especially in 
moist or wet habitats. Preliminary sampling 
in Peninsular Florida (unpublished) indi- 
cates that the same is true there. The genus 
is mainly tropical; only one species is re- 
corded from Europe and is uncommon. Eu- 
rope, India, Japan, Africa, South America. 

Hemienchytraeus stephensoni 

(Cognetti, 1927) 

Fig. 5, A-I 

Enchytraeus cavicola Stephensen, 1924 
(non Joseph 1880):127-129, PI. 6, figs. 
\^. 

Enchytraeus stephensoni Cognetti, 1927:4. 

Enchytraeus myrmecophilus Cernosvitov, 
1930a: 85-89, figs. 1-9. 

Enchytraeus rangoonensis Stephensen, 
1931:177-179, fig. 1. 

Hemienchytraeus stephensoni (E. cavicola 
— E. stephensoni + E. myrmecophilus + 
E. rangoonensis) Cernosvitov, 1934:298- 
304, figs. 1-11. 

Hemienchytraeus stephensoni Cernosvitov 
1939:92-93, figs. 43^9; Christoffersen 
1979:40-46, figs. 1-23; Nakamura 1984: 
32-33, Fig. IB, C, F; Ddzsa-Farkas 
1989:200-202, figs. 24-33. 

Material examined. — Five stained, whole 
mounts, USNM 170741-170745, four 
stained, whole mounts ROMIZ 13239- 
13240, 13246-13247; 30 stained, whole 
mounts in the author's collection. Approx- 
imately 50 live specimens examined. 

Description of new material. — Small to 



VOLUME 109, NUMBER 1 



131 




Fig. 5. Hemienchytraeus stephensoni. A, peptonephridium; B, sperm funnel, live, worm; C, spermatheca; 
D, preclitellar nephridium; E, postclitellar nephridium; F, brain; G, anterior setal bundle; H, posterior setal bundle; 
I, coelomocytes. Hemienchytraeus bifurcatus. J, peptonephridium, K, sperm funnel, live worm; L, spermatheca. 
Buchholzia fallax. M, esophageal gland in IV. pb primary branch, sb secondary branch, tb tertiary branch. 



medium-sized worms, live specimens rather 
transparent, with a prominent clitellar re- 
gion due to presence of large eggs. Live 
specimens 8-10 mm, fixed specimens 6-9 
mm, diameter 0.2-0.3 mm, 0.24-0.38 mm 
at cHtellum. Segments 36-48 {X = 44. 1 , SD 
= 2.7, n = 29). Setae two per bundle, oc- 
casionally three, with weak ental hook. Se- 
tae absent in XII. In anterior region setae 
straight or slightly curved and rather small, 
32^0 |jim long, more curved, thicker and 
longer in posterior half of worm, 50-62 iJtm 
in terminal segments (Fig. 5G, H). Cuta- 
neous gland cells small and inconspicuous, 
in about four rows per segment. Prostomi- 
um with numerous dark-staining gland 
cells. Clitellum over XII-y2XIII, gland cells 
small, arranged in more or less regular 
transverse rows, especially in ventral region 
and on anterior and posterior borders of cli- 
tellum. Glands may be absent mid-ventral- 
ly. Head pore near tip of prostomium. Some 



specimens were in mucus tubes encrusted 
with particles of organic matter. 

Three pairs of pharyngeal glands, all 
united dorsally and with ventral lobes. Pep- 
tonephridium arises from dorsal side of 
pharynx in III, proximal portion of variable 
length, thick-walled with a coiled or sinu- 
ous lumen, primary and secondary branch- 
es narrower but also with coiled lumina 
(Fig. 5A). Each secondary branch divides 
into three to five thin-walled, finger-like 
processes. Entire peptonephridium confined 
to IV. Chloragocytes from V, form a dense 
layer from VII, 11-13 cells across intestine 
in preclitellar region in compressed speci- 
mens. Transition between esophagus and 
intestine gradual with enlargement behind 
pharyngeal glands. In all specimens col- 
lected from the Pensacola region in 1993, a 
section of the intestine behind the clitellum, 
usually between XIV-XVI and XXVIII- 
XXX, was enlarged and brown in colour. In 



132 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Other specimens, the brown region was re- 
duced or absent. Coelomocytes sparse, of 
various sizes and shapes, but mainly round, 
8-18 |jLm, almost hyaline in live individu- 
als, faintly granular with a distinct nucleus 
in fixed specimens (Fig. 51). Dorsal vessel 
originates in XIII, usually with isolated, 
yellow, dendritic cells on surface of vessel 
(seen in live worms); anterior bifurcation at 
0/1. Blood colorless. Nephridia with large, 
ovoid preseptale that encloses a dorsally di- 
rected nephrostome, postseptale of variable 
shape, usually ovoid in preclitellar seg- 
ments where duct arises mid-ventrally and 
elongated in posterior segments where duct 
is sub-terminal or terminal (Fig. 5D, E). 
First nephridium at 6/7. Brain about 1.5 
times longer than wide, deeply indented or 
even cleft on its anterior border, posterior 
border sinuous or straight (Fig. 5F). 

Testes compact. Small, paired seminal 
vesicles usually present in XI. Sperm fun- 
nels funnel-shaped, somewhat flattened, 
190-360 |jLm long, 40-1 12 |jim wide, some- 
what longer than diameter of worm, gen- 
erally six to eight times longer than wide in 
live specimens, with collar about same 
width or slightly narrower than rest of fun- 
nel (Fig. 5B). Funnels taper to spirally 
coiled ducts 5-6 |xm in diameter, which 
open at small, compact penial bulbs, lon- 
gitudinal diameter 54-82 ixm. Male pores 
curved slits. One to three large eggs present 
at a time, contained in an egg sac that ex- 
tends to 14/15 or rarely 15/16. Spermathe- 
cae with long, thick-walled ectal ducts, 12- 
18 |xm in diameter, outer surface rough, 
which swell to form sperm-containing 
chambers in V or VI, continue as smooth, 
thin-walled tubes, and terminate in thin- 
walled, cylindrical ampullae in VI-IX (Fig. 
5C). Length of spermathecae variable; am- 
pullae may be twice or several times longer 
than wide, swollen and occupying up to IVi 
segments when mature. 

Remarks. — Hemienchytraeus stephen- 
soni is a very variable species, hence the 
number of nominal species synonymised by 
Cernosvitov (1934). The variability has 



been commented on by Cernosvitov (1939), 
who noted differences in the length of the 
basal part of the peptonephridium and in the 
form and complexity of the septal (pharyn- 
geal) glands in South American specimens. 
Christoffersen (1979) distinguished four 
types among Brazilian material from a sin- 
gle locality that differed in body length and 
width, setal length, pharyngeal glands, form 
of the peptonephridium, origin of dorsal 
vessel, number and shape of nephridia, 
form of the penial bulbs, length of sperm 
funnels and length of spermathecae and am- 
pullae. He concluded that H. stephensoni is 
a complex of cryptic species. Two extreme 
forms were recognised among West Florida 
material that differed in lengths of setae, 
sperm funnels, spermathecal ampullae and 
the proximal section of the peptonephri- 
dium and in the form of the pharyngeal 
glands, but intermediates were also found. 
Size and segment number and dimensions 
of setae and sperm funnel are within the 
range of specimens from South America 
(Christoffersen 1979, Dozsa-Farkas 1989) 
and Japan (Nakamura 1984), but specimens 
differ in having both secondary and tertiary 
branches to the peptonephridium and in the 
presence of seminal vesicles. No individu- 
als were found with more than two second- 
ary branches on each side of the peptone- 
phridium, as in material from Lake Titicaca 
(Cernosvitov 1939) and Ecuador (Dozsa- 
Farkas 1989). In spite of Christoffersen's 
ability to recognise distinct types within a 
region, and the differences between mate- 
rial from Florida and South America, I be- 
lieve H. stephensoni to be one variable spe- 
cies with a wide ecological tolerance and a 
wide geographic range. 

Habitat and distribution. — Hemienchy- 
traeus. stephensoni was by far the most 
common enchytraeid in the region and was 
recorded from all localities and most kinds 
of inland habitat. 

Pensacola, woodland, swamp, edge of a 
pond, floodbank of a river and boggy area, 
Avalon Peninsula, pitcher plant (Sarracen- 
ia) bog; Ponce de Leon, woodland at edge 



VOLUME 109, NUMBER 1 



133 



of a stream; Lake wood, edge of a lake; 
Marianna, woodland on limestone and bank 
of a river; Torreya State park, woodland on 
limestone; Port St. Joseph, edge of roadside 
ditch. pH range 3.6-7.2. 

Mainly tropical. South America (Argen- 
tina, Paraguay, Brazil, Bolivia, Ecuador), 
India, Burma, Central Africa, Japan. Re- 
corded from almost every conceivable ter- 
restrial habitat including woodland, grass- 
land, tree bark, anthills, bat guano and the 
marine littoral zone. New for North Amer- 
ica. 

Hemienchytraeus bifurcatus 

Nielsen & Christensen, 1959 

Fig. 5J-L 

Hemienchytraeus bifurcatus Nielsen & 
Christensen 1959:45, figs. 23-27. 

Hemienchytraeus bifurcatus Nakamura 
1984:31-32, fig. lA, D, E.; Kasprzak 
1986:178, figs. 639-643. 

Material examined. — Three stained, 
whole mounted specimens USNM 170746- 
170748; one stained whole mount ROMIZ 
13248; seven stained whole mounts in the 
author's collection. Approximately 28 live 
specimens examined. 

Description of new material. — Medium 
to small worms, live specimens 4-6 mm, 
transparent when viewed microscopically, 
with a prominent clitellar region due to 
presence of rather large eggs. Fixed length 
4-6 mm, diameter 0.3-0.4 mm, 0.3-0.45 
mm at clitellum. Segments (25)29-44 (X = 
35.2, SD = 4.9, n = 20). Setae straight or 
slightly curved with a weak ental hook, two 
per bundle throughout, absent in XII, 32- 
58 ixm. Cutaneous glands small, indistinct. 
Clitellum over Xll-i/aXIII with small gland 
cells arranged in more or less transverse 
rows or sometimes irregularly, smaller or 
absent in mid-ventral region. Head pore 
near tip of prostomium. Some specimens 
were in mucus tubes with adhering debris 
and soil. 

Three pairs of pharyngeal glands, all 
lobed and united dorsally. Peptonephridium 



arises from mid dorsal region of pharynx in 
III, proximal section of variable length, 
branching dichotomously into two primary 
and four secondary branches (Fig. 5J). 
Proximal part of unpaired section hollow 
and contractile in live specimens, distal sec- 
tion and primary branches stout with a 
coiled or sinuous lumen, secondary branch- 
es thin-walled with a wide lumen. Entire 
peptonephridium confined to IV. Chlora- 
gocytes from V, forming a dense layer from 
VII. Esophageal-intestinal transition gradu- 
al, gut expands behind pharyngeal glands. 
Coelomocytes sparse, of various shapes and 
sizes, dominant type round or oval, 10-14 
|jLm, nucleated, granular. Dorsal vessel orig- 
inates in XIII or XIV, anterior bifurcation 
at 0/1. Blood colorless. Nephridial presep- 
tale large, postseptale egg-shaped with pos- 
tero-ventral efferent duct in preclitellar re- 
gion, more elongated with terminal or sub- 
terminal duct behind clitellum. Nephridia 
starting at 4/5 or 5/6, absent X-XVI. Brain 
rectangular, about 1.5 times longer than 
wide, deeply indented or even cleft on its 
anterior border, more or less truncated pos- 
teriorly. 

Small, paired seminal vesicles usually 
present, sometimes only developed on one 
side. Sperm funnels funnel-shaped, three or 
four times longer than broad in live worms, 
100-150 |xm in length, equal to Vs or Vz di- 
ameter of worm, with a collar equal to or 
slightly narrower than rest of funnel (Fig. 
5K). Funnels taper to long, spirally-coiled 
ducts in XII, open at small, compact penial 
bulbs, about 32 [xm across. One to three 
large eggs present at a time, egg sac ex- 
tending to XIV. Spermathecae long, each 
consists of a fairly stout, thick-walled ectal 
duct, without glands at the orifice, duct 
swells to form a sperm-containing chamber 
in V or VI and then extends as a thin-walled 
tube to an ovoid or spherical ampulla situ- 
ated in any of segments VI-IX, usually in 
VII (Fig. 5L). 

Remarks. — This species, like the previ- 
ous one, is very variable, especially in size, 
form of the nephridia and lengths of sper- 



134 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



mathecae and sperm funnels. The different 
dimensions of the reproductive organs are 
not related to stage of maturity, for individ- 
uals with sperm in the ampulla may have 
long or short spermathecae and either short 
or long sperm funnels. Danish specimens 
(Nielsen & Christensen 1959) differed in 
having a brain indented posteriorly, ne- 
phridia with terminal efferent ducts, coe- 
lomocytes with refringent granules, no sem- 
inal vesicle and a sperm funnel only 2-3 
times longer than wide. A longer sperm 
funnel was also reported in Japanese spec- 
imens (Nakamura 1984). 

The species is similar in general appear- 
ance to H. stephensoni but is generally 
smaller and has somewhat more granular 
coelomocytes, more noticeable in live spec- 
imens. The two species often occurred to- 
gether and although the form of the pepto- 
nephridium is quite different, this was 
sometimes difficult to see clearly. Mature 
individuals could be reliably distinguished 
by the greater dimensions of the sperma- 
thecal ampulla and sperm funnel in H. ste- 
phensoni, as noted by Nakamura (1984). 

Habitat and distribution. — Not as abun- 
dant, nor as widespread as H. stephensoni. 
Pensacola, woodland leaf litter at the Uni- 
versity of West Florida, grass cuttings at 
Scenic Hills golf course; Avalon Peninsula, 
pitcher plant (Sarracenia) bog; Ponce de 
Leon, woodland soil and leaf litter; Marian- 
na, woodland soil and leaf litter on lime- 
stone. pH range 3.9-6.9. Denmark, France, 
Japan, India. New for North America. 

Genus Henlea Michaelsen, 1889 

The principal diagnostic characters of the 
genus are straight or slightly curved setae, 
usually of unequal size within a bundle, the 
outer ones longer, esophagus expands 
abruptly into the intestine (with the possible 
exception of the sub-genus Hepatogaster), 
esophageal appendages present, intestinal 
diverticula present or absent, a preclitellar 
origin of the dorsal vessel, usually in VIII 
or IX, and nephridia with free nephrostome 



and anterior origin of the efferent duct, of- 
ten described as ''Henlea type" but not 
unique to that genus. 

Cosmopolitan: E and W of North Amer- 
ica, S America, Europe, Siberia, Africa, 
Antarctic. 



Henlea perpusilla Friend, 1911 
aug. Cernosvitov 1937b 

Henlea perpusilla Friend, 1911:466^67. 

Henleanella perpusilla Friend, 1913a:89. 

Henlea bisetosa Friend, 1914:135. 

Henlea inusitata Friend, 1913a:83-84. 

Henlea minima Friend, 191 3a: 84. 

Henlea nivea Cernosvitov, 1930b:88, figs. 
3-6. 

Henlea perpusilla Friend 1913b:270-271, 
figs. 34-35; Cernosvitov 1937a: 194-196, 
figs. 2-5; Nielsen & Christensen 1959: 
58-59, figs. 46, 47; Kasprzak 1986:256- 
257, figs. 992-996. 

Material examined. — Two stained, whole 
mounts USNM 170749-170750; two 
stained whole mounts in the author's col- 
lection. Live specimens examined, 27. 

Remarks. — The large number of syn- 
onyms that are now recognised reflects a 
wide variation in size and morphology 
(Cernosvitov 1941, Nielsen & Christensen 
1959), which is partly explained by the ex- 
istence of different cytotypes (Nielsen & 
Christensen 1959). Size of the live speci- 
mens (6-8 mm) and segment number (30- 
33) of Florida material are within the range 
of European specimens. 

Habitat and distribution. — Common in 
Marianna State Park, deciduous woodland 
on limestone, sandy humus, sandy alluvial 
deposits and humus with limestone frag- 
ments; Tall Timbers Research Station, live 
oak forest, dark, sandy loam and leaf litter. 
pH range 5.5-7.1. Absent from the west of 
the study area. Canada (Cornwallis Is., 
Prince of Wales Is., Devon Is., Rocky 
Mountains), Greenland, Europe, Lebanon, 
Siberia, Japan, Antarctic, Bolivia. 



VOLUME 109, NUMBER 1 



135 



Henlea ventriculosa 
d'Udekem, 1854 

Enchytraeus ventriculosus d'Udekem, 1854: 
863, figs. 1, 4, 6-9. 

Henlea multispinosa Friend, 1913a:85. 

H. (Udekemiana) ventriculosa Cernosvitov, 
1930b:75. 

Henlea groenlandica Cernosvitov, 1929: 
146, figs. 1, 2. 

H. {Udekemiana) groenlandica Cernosvi- 
tov, 1930b:75. 

Fridericia steward Stephenson, 1909:109. 

Henlea ventriculosa Michaelsen, 1889:31- 
32, 1900:69-70; Nielsen & Christensen 
1959:62, figs. 50, 53, 57; Kasprzak 1986: 
264-265, figs. 1028-1030. 

Material examined. — Two stained, whole 
mounts USNM 170751-170752; three 
stained, whole mounts in the author's col- 
lection. Five live specimens examined. 

Remarks. — Size, 5-8 mm when fixed, 
and segment number (38-44) of Florida in- 
dividuals are within the range of European 
specimens but the spermathecae have short- 
er ectal ducts, about equal to the length of 
the ampullae, while each ental duct is lon- 
ger than the combined lengths of ampulla 
and ectal duct. In addition, there are several 
small glands at the ectal opening. Danish 
specimens lacked these glands, the ectal 
duct was longer and the ental duct shorter 
(Nielsen & Christensen 1959). In spite of 
these differences, the species could be iden- 
tified with confidence by the characteristic 
gut diverticula. 

Habitat and distribution. — Marianna 
State Park, deciduous woodland on lime- 
stone, rotting wood, leaf litter, humus with 
limestone fragments. pH range 5.3-6.7. 
Canada (Cornwallis Is., Prince of Wales 
Is.), USA (Massachusetts, N. Carolina), Eu- 
rope, Siberia, Tibet, Japan, New Zealand, S. 
America. 

Genus Buchholzia Michaelsen, 1887 

The genus is recognised by the following 
unique combination of characters: sigmoid 



setae, decreasing in size within bundles to- 
ward the dorsal and ventral midlines of the 
body; hollow esophageal appendages in IV; 
transition between esophagus and intestine 
abrupt with one or two dorsal diverticula at 
7/8; anteclitellar origin of the dorsal vessel 
in the region of the intestinal diverticula; 
and small, hyaline, anucleate coelomocytes 
in addition to the normal kind. 

So far, confined to the northern hemi- 
sphere but not recorded from Asia. Canada, 
Europe, N. Africa. 

Buchholzia fallax Michaelsen, 1887 
(Fig. 5M) 

Buchholzia fallax Michaelsen, 1887:374- 
376, pi. 21, fig. 4A-E. 

Buchholzia fallax Beddard, 1895:334-335; 
Michaelsen 1900:72-73; Cernosvitov 
1928:9; Ude 1929:57-58; Nielsen & 
Christensen 1959:65, figs. 59-61; Wilcke 
1967:72, fig. 6A, B; Kasprzak 1986:117- 
118, figs. 294-298, table 8. 

Material examined. — Two stained, whole 
mounts USNM 170753-170754; ten whole 
mounts in the author's collection. Live 
specimens examined, 18. 

Remarks. — Distinguished from B. appen- 
diculata (Buchholz, 1862), which was re- 
corded from Montreal by Nurminen 
(1973b), by the presence of one, rather than 
two, intestinal diverticula and only three in- 
stead of four pairs of pharyngeal glands, 
and by having male organs in the usual po- 
sition for enchytraeids. All specimens ex- 
amined are small, 4-7 mm compared with 
12-14 mm for specimens from Denmark 
(Nielsen & Christensen 1959). Segments 
30-38 (Danish material 37-42). A seminal 
vesicle is usually present, stated to be ab- 
sent by Nielsen & Christensen and not men- 
tioned by Michaelsen (1887). The pulsating 
esophageal appendages in IV are of a dif- 
ferent shape from those figured by Nielsen 
& Christensen (1959:143, fig. 60); they are 
elongate with partial cross walls dividing 
the cavity into three compartments (Fig. 
5M), rather than rosette-like. The small. 



136 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



secondary cavities in the spermathecal am- 
pulla, figured by Michaelsen, could not be 
seen and have not been mentioned by other 
authors. Some specimens were inside mu- 
cus tubes encrusted with rings of soil par- 
ticles. 

Habitat and distribution. — Common in 
Marianna State Park, deciduous woodland 
on limestone. A few in a similar habitat in 
Torreya State Park. pH range 5.5-7.1. Ab- 
sent from the west of the study area. Eu- 
rope, N. Africa. New for North America. 

Acknowledgments 

Support for this work was provided by 
the Mary Ball Washington Foundation at 
the University of West Florida, Pensacola. 
I am grateful to members of the Department 
of Biology for useful information on the re- 
gion and in particular to Paul Hamilton for 
his help in locating the wide range of hab- 
itats sampled. The friendly hospitality of 
the Department was much appreciated. 

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15:1-49, plates 1-16. 



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chytraeidae during expeditions. — Soil Biology 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1): 138-142. 1996 

Artacama valparaisiensis, a new species of Terebellidae 

(Annelida: Polychaeta) from sub tidal soft bottoms of 

Valparaiso Bay, Chile 

Nicolas Rozbaczylo and Marco A. Mendez 

Departamento de Ecologia, Facultad de Ciencias Biologicas, P. Universidad Catolica de Chile, 

Casilla 114-D, Santiago, Chile 

Abstract. — A new species of terebellid, Artacama valparaisiensis, is de- 
scribed from Valparaiso Bay, Central Chile. The species closely resembles A. 
crassa Hartman, 1967, from the South Shetland Islands, Antarctica, but can be 
distinguished on the basis of the following morphological criteria: shape and 
number of proboscidial papillae, size and number of the branchial filaments, 
relative size of lobe at the upper edge of uncinal ridges in abdominal parapodia 
and presence of constrictions in the limbate setae. 



Systematics of the Terebellidae was re- 
viewed recently by Holthe (1986). Accord- 
ing to his review, Artacama Malmgren, 
1866, with eight valid species is the only 
genus in subfamily Artacaminae. McHugh 
(1995) has carried out a cladistic analysis 
of the Terebellidae in which she shows that 
Artacama is the sister taxon of a large clade 
within the Amphitritinae. The Artacaminae 
has therefore been synonymized with the 
Amphitritinae, which is diagnosed by the 
presence of double rows of uncini on pos- 
terior thoracic segments. The character that 
identified the Artacaminae, a peristomium 
modified on its ventral side to form a con- 
spicuous proboscis-like organ adorned with 
papillae, is considered an autapomorphy for 
the genus Artacama (McHugh 1995). Only 
two species of Artacama, A. crassa Hart- 
man, 1967 and A. proboscidea Malmgren, 
1 866, have been reported from southeastern 
Pacific, in antarctic and subantarctic waters 
(Rozbaczylo 1985). From 1978 to 1980 an 
extensive research program on benthic 
communities at Valparaiso Bay, Central 
Chile, was carried out by Dr. Jose Stuardo 
(presently at Universidad de Concepcion) 
and Dr. Hector Andrade (presently at A & 
A Tecnolab S.A., Division Ambiental). 



Three areas with different particle size 
(sand, sandy-mud and sandy-silt), organic 
matter and other biochemical components 
were sampled, from 34 to 59 meters depth. 
Samples from these areas have been studied 
by Stuardo et al. (1981) regarding their bio- 
chemical and granulometric composition. 
They showed that among the three sampled 
areas there were differences in proteins, lip- 
ids and organic matter; the sandy-mud area 
showed the highest values and sandy area 
the lowest values. Among the abundant 
polychaetes collected during that program 
numerous individuals of Artacama were 
found. These differed from the other known 
species reported for the genus, and conse- 
quently are described as a new species. 

Specimens of the new species were com- 
pared with the holotype of A. crassa de- 
posited in the National Museum of Natural 
History, Washington, D.C. (USNM 55569) 
and a specimen of A. proboscidea from 
Western Canadian Arctic (USNM 41132) 
identified by E. & C. Berkeley. 

Type and paratype specimens of the new 
species are deposited in the National Mu- 
seum of Natural History, Smithsonian In- 
stitution, Washington, D.C. (USNM); Sala 
de Sistematica, Departamento de Ecologia, 



VOLUME 109, NUMBER 1 



139 



Pontificia Universidad Catolica de Chile, 
Santiago (SSUC); and Museo de Zoologia, 
Universidad de Concepcion (UCCC). 

Materials and Methods 

Polychaetes were collected at Valparaiso 
Bay, Central Chile. Samples were taken 
monthly, from November 1979 to October 
1980, with a 0.1 m- Van Veen dredge at 
three areas with different particle size 
(sand, sandy-mud and sandy-silt), from 34 
to 59 meters depth. Polychaetes were fixed 
in 10% formalin and preserved in 70% eth- 
anol. Figures were prepared by means of a 
drawing tube on a Wild M-5 microscope. 
Scanning electron microscopic (SEM) ob- 
servations and photographs were obtained 
using a JEOL JSM-25SII microscope. 

Artacama valparaisiensis, 

new species 

Figs. 1, 2 

Material examined. — Central Chile: Val- 
paraiso Bay, off Punta Osas, ca. 32°59'20"S, 
71°33'56"W, 44 m, H. Andrade, coll., 9 Jul 
1979, holotype (USNM 170005), and 14 
paratypes (USNM 170006); 4 Sep 1979, 8 
paratypes (UCCC 23169-23176), and 14 
paratypes (SSUC 6719). 

Description. — Holotype incomplete with 
52 setigers, measuring about 60 mm long, 
width about 9 mm at thorax. Body (Fig. la) 
thick anteriorly and colorless in alcohol, 
with first 8-11 segments enlarged, tapers 
posteriorly to a narrow pygidial end. Total 
length of one of the largest complete spec- 
imens, is 104 mm including proboscis, 
width 12 mm at thorax and 4 mm at ab- 
domen, and consists of 102 setigerous seg- 
ments; 17 thoracic setigers and 85 abdom- 
inal setigers. Large, ovoid and papillose 
proboscis (Fig. la) extends forward from 
buccal segment below mouth. Prostomium 
(Fig. lb) is small folded, bilobed structure, 
with small oral aperture immediately below. 
Eyes absent. Small, horseshoe-shaped ten- 
tacular lobe on dorsal side of peristomium, 
with a dorsal indentation and numerous ten- 



tacles, short and clubbed, most of which are 
broken. Proboscis covered overall with nu- 
merous, conical and minute, papillae (Fig. 
Ic) arranged in about ninety longitudinal 
rows. Lateral lappets absent on segments 2- 
4. Three pairs of long filiform branchiae on 
segments 2—4; each branchia is a tuft of ap- 
proximately 50 equal-length filaments aris- 
ing from basal stump. Nephridiopores, in 
form of short tubes, number five pairs; the 
best developed are on segment 3 below sec- 
ond pair of branchiae, in line with the more 
posterior notopodia; smaller ones are on 
segments 6, 7, 8 and 9, between notopodia 
and uncinal ridges, postero-ventral to no- 
topodia. Ventral glandular pads present on 
first 10 setigers. Thorax with fixed number 
of setigers; seventeen bundles of notosetae 
starting on segment 4; uncini first present 
from the second setiger (fifth segment); oc- 
cur in single rows on first six uncinal ridg- 
es, then in double rows from setigers 8 to 
17, oriented "fang to fang." Thoracic no- 
topodia (Fig. Id) with triangular lamellae, 
postsetal lobe larger than presetal; 40-50 
long, pointed setae decreasing in length 
from dorsal to ventral part of notopodia, lat- 
erally winged, with one or two constrictions 
(Fig. le). Thoracic neuropodia with avicu- 
lar uncini with a large fang surmounted by 
four or five alternating rows of many small 
teeth (Fig. 2a, b). Abdomen with numerous 
segments bearing parapodia (Fig. Ig, h) 
with flaplike tori and dorsally large, subcir- 
cular membrane which increase in size to- 
wards the posterior end; uncini in single 
rows restricted to ventral margin; abdomi- 
nal uncini resemble thoracic but main fang 
is longer and thinner and have six or seven 
alternating rows of teeth (Figs. If, 2c, d). 

Posterior end tapers to terminal pygidium 
(Fig. li), with terminal anus, with crenulate 
sides. 

Geographical distribution. — Known only 
from Valparaiso Bay, Chile. 

Remarks. — Artacama valparaisiensis 
new species resembles, most closely, A. 
crassa Hartman, 1967 and A. proboscidea 
Malmgren, 1866. However A. valparaisien- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. L Artacama valparaisiensis new species (USNM 170005). a, entire animal in left lateral view; b, 
prostomium in dorsal view with the oral aperture immediately below; c, proboscidial papillae; d, thoracic par- 
apodium, right side anterior view; e, limbate thoracic notoseta from fifth setiger, in lateral view showing con- 
strictions; f, abdominal uncinus in frontal and lateral view, from tenth abdominal parapodium; g, second abdom- 
inal parapodium, right side anterior view, with large foliaceous dorsal lobe and uncinal torus; h, tenth abdominal 
parapodium, right side anterior view, with large foliaceous dorsal lobe and uncinal torus; i, posterior end, in 
lateral view. Scales = 1 cm for a; 1 mm for b, g, h, i; 0.5 mm for c, d; 0.01 mm for f; 0.1 mm for e. 



VOLUME 109, NUMBER 1 



141 




Fig. 2. Artacama valparaisiensis new species, SEM micrographs: a, thoracic uncini in lateral view from the 
first uncinigerous segment (fifth thoracic segment); b, detail of the upper part of an uncinus of thoracic segment 
1 1 (first segment with double rows of uncini); c, abdominal uncini in lateral view from the fourth abdominal 
segment; d, detail of the upper part of an uncinus of the fourth abdominal segment. Scales = 0.01 mm for a-d. 



sis differs from A. crassa in that papillae of 
the proboscis are more abundant, conical, 
and smaller; the number of branchial fila- 
ments is higher (ca. 50 in each branchiae) 
and each filament is longer and thinner; 
presence of constrictions in the limbate se- 
tae; thoracic notopodia with postsetal la- 
mellae longer than presetal lobe, while in 
A. crassa both lobes are short; abdominal 
parapodia with the uncinigerous tori longer 
and the dorsal membrane comparatively 
smaller. On the other hand, A. proboscidea 
shows fewer branchial filaments, shorter 
and thicker than those of A. valparaisiensis. 



Papillae of the proboscis of A. proboscidea 
are bigger and fewer than in A. valparais- 
iensis. Dorsal membrane of abdominal par- 
apodia of A. valparaisiensis is bigger than 
those of A. proboscidea. Absence of con- 
strictions in the limbate setae of A. probos- 
cidea. Mcintosh (1885) described A. chal- 
lengeriae from several localities off Ker- 
guelen Islands, 46-202 m; Hartman (1959) 
considers this species similar to A. probos- 
cidea. Description of A. challengeriae is so 
general and vague that it is difficult to com- 
pare it with A. valparaisiensis. Neverthe- 
less, from figures (PI. LI. fig. 6; PI. 



142 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



XXVIIIA. figs. 23a, b, 24, 25) in Mcintosh 
(1885) can be established that A. challen- 
geriae differs from A. valparaisiensis at 
least in that the former has proboscidial pa- 
pillae bigger an fewer; edges of limbate se- 
tae are entire, without constrictions; ventral 
margin of uncini presents an ondulating 
outline while in A. valparaisiensis has a 
uniform and slight convexity the same as in 
A. crassa and A. proboscidea. 

Etymology. — The specific name valpa- 
raisiensis refers to the type locality, Val- 
paraiso Bay. 

Ecological remarks. — Artacama valpa- 
raisiensis was more abundant between May 
and September, with densities of 100 indiv./ 
m^ in May and 250 indiv./m^ in September. 
The species showed the highest density in 
the sandy-mud area (30-250 indiv./m-), the 
second one was the sandy-silt area (5-100 
indiv./m^); at the sandy area the species was 
extremely rare. In the sandy area few spec- 
imens of small size were found only in May 
and September while in the sandy-mud and 
sandy-silt areas numerous specimens of dif- 
ferent size were found throughout the sam- 
ple period. This pattern of distribution ap- 
pears to be correlated with the different or- 
ganic matter content of sediments in the 
sites sampled (Stuardo et al. 1981). In the 
population studied it was possible to distin- 
guish adult females by the presence of eggs 
in the coelomic cavity. Maximum egg size 
was 165 |xm diameter. 

Acknowledgments 

We thank Dr. Jose Stuardo, Departamen- 
to de Oceanologia, Universidad de Concep- 
cion, who made specimens available for 
sudy and Dr. Hector Andrade, A & A Tec- 
nolab S.A., Division Ambiental who fur- 
nished information about collecting. We 
specially thank Clara Yanez for her skill 
and patience in producing the illustrations 



used in this paper and Jose Morillas for the 
scanning electron photomicrographs. We 
thank Dr. Kristian Fauchald of the National 
Museum of Natural History, Washington, 
D.C., for the loan of specimens for com- 
parison. The manuscript was greatly im- 
proved by the valuable comments and sug- 
gestions provided by Dr. Christopher J. 
Glasby and an anonymous reviewer who 
also provided valuable bibliography. 

Literature Cited 

Hartman, O. 1959. Catalogue of the polychaetous an- 
nelids of the World, part II. — Allan Hancock 
Foundation Publications Occasional Paper 23: 
355-628. 

. 1967. Polychaeta annelids collected by the 

USNS Eltanin and Staten Island Cruises, chief- 
ly from Antarctic Seas. — Allan Hancock Mono- 
graphs in Marine Biology 2:1-387. 

Holthe, T. 1986. Evolution, systematics, and distri- 
bution of the Polychaeta Terebellomorpha, with 
a catalogue of the taxa and a bibliography. — 
Gunneria 55:1-236. 

Malmgren, A. J. 1866. Nordiska Hafs-Anunlater. — 
Ofversigt af Konglia Vetenskaps-Akademiens 
Forhandlingar, Stockholm 22:355^10, pis. 18- 
29. 

McHugh, D. 1995. Phylogenetic analysis of the Am- 
phitritinae (Polychaeta: Terebellidae). — Zoolog- 
ical Journal of the Linnean Society (in press). 

Mcintosh, W C. 1885. Report on the Annelida Poly- 
chaeta collected by H.M.S. Challenger during 
the Years 1873-1876. — In Report on the Sci- 
entific Results of the Voyage of H.M.S. Chal- 
lenger during the Years 1873 under the Com- 
mand of Captain George S. Nares, R.N., F.R.S. 
and the Late Captain Frank Tourle Thomson, 
R.N., Zoology, 12(34): 1-554, plates 1-55, and 
1A-39A, 1 map. 

Rozbaczylo, N. 1985. Los Anelidos Poliquetos de 
Chile. Indice sinonimico y distribucion geo- 
grafica de especies. — Monografias Bioldgicas 
3:1-284. 

Stuardo, J., M. A. Soto, H. Andrade, & R. Aguilar. 
1981. Caracteristicas granulometricas y com- 
ponentes bioqufmicos de los sedimentos de tres 
estaciones submareales de Valparaiso. — ^Revista 
de Biologia Marina, Instituto de Oceanologia, 
Universidad de Valparaiso, 17(2): 17 1-196. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1):143-149. 1996 

Review of Hermilepidonotus Uschakov, 1974, and two species of 
polynoid polychaetes (Lepidonotinae) 

Marian H. Pettibone 

Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — The genus Hermilepidonotus Uschakov, 1974, and type species 
Lepidonotus robustus Moore, 1905, are reviewed, and compared with the close- 
ly related Lepidonotus helotypus (Grube, 1877), based on examination of types 
and available new material. 



Under Lepidonotus helotypus (Grube, 
1877), from China, Seidler (1924:56) in- 
cluded Lepidonotus robustus Moore, 1905, 
from the Gulf of Alaska. This was followed 
by Annekova (1937), from the North Japan 
Sea, by Hartman (1938), from Alaska, and 
by Uschakov (1950, 1955), from the 
Okhotsk Sea. On further study, L. robustus 
was considered to be distinct from L. hel- 
otypus, by Hartman (1948) and by Uschak- 
ov (1974). Based mainly on a characteristic 
pharynx, the latter author referred L. ro- 
bustus to the new genus Hermilepidonotus. 
The close similarity of these two Pacific 
Ocean boreal species is the basis for adding 
L. helotypus to Hermilepidonotus. The two 
species are more fully described herein and 
some new records are presented. In addition 
to the specimens deposited in the National 
Museum of Natural History, Smithsonian 
Institution (USNM), types and additional 
material were obtained on loan or in ex- 
change from the following sources: the for- 
mer British Museum (Natural History), 
London (BMNH), now renamed the Natural 
History Museum, London, through J. D. 
George and A. I. Muir; National Science 
Museum, Tokyo (NSMT), through M. Ima- 
jima; Zoological Institute Academy of Sci- 
ence, Leningrad (ZIASL), through P. V. Us- 
chakov; Zoologisches Museum, Berlin 
(ZMB), through G. Hartwich; and Zoolo- 
gisches Staatsmuseum, Hamburg, through 
G. Hartmann-Schroder. 



Family Polynoidae Kinberg, 1856 

Subfamily Lepidonotinae Willey, 1902 

Genus Hermilepidonotus Uschakov, 1974, 

emended 

Type species. — Lepidonotus robustus 
Moore, 1905, by monotypy and original 
designation. 

For: H. robustus (Moore, 1905) and H. 
helotypus (Grube, 1877), new combination. 

Diagnosis. — Body elongate-oval, with 
26 segments. Prostomium lepidonotoid, bi- 
lobed, with 3 antennae and 2 palps; cera- 
tophore of median antenna in anterior 
notch, lateral antennae inserted terminally 
on anterior extensions of prostomium, on 
same level as median antenna; 2 pairs of 
eyes on posterior half of prostomium. First 
or tentacular segment not distinct dorsally; 
tentaculophores lateral to prostomium, each 
with aciculum, few setae on inner side, and 
pair of dorsal and ventral tentacular cirri, 
similar to antennae. Elytra and large ely- 
trophores 12 pairs, on segments 2, 4, 5, 7, 
9, 11, 13, 15, 17, 19, 21, 23. Elytra large, 
suboval, leaving middorsum uncovered, 
without fringes of papillae, thick, soft, with 
micropapillae on {H. robustus) or with mi- 
crotubercles and conical macropapillae (//. 
helotpyus). Dorsal cirri on non-elytrigerous 
segments, with cylindrical cirrophores and 
smooth styles; dorsal tubercles bulbous in- 
flated areas. Second or buccal segment with 
first pair of elytrophores, biramous para- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



podia, and long ventral buccal cirri lateral 
to ventral mouth. Eversible muscular phar- 
ynx with 2 pairs of chitinous hooked jaws 
and border papillae; [about 15 pairs (13-17, 
in H. helotypus) or border papillae plus nu- 
merous branched, threadlike papillae, in H. 
robustus)]. Parapodia biramous, with small 
conical notopodia on anterodorsal sides of 
large neuropodia; neuropodia with subequal 
presetal and postsetal lobes, with smaller, 
truncate supraacicular parts and longer, di- 
agonally truncate subacicular parts. Noto- 
setae slender, with spinous rows, short, with 
blunt tips and longer, with fine tips. Neu- 
rosetae stout, with long spinous regions and 
long, bare, entire tips. Ventral cirri short, 
subulate. Nephridial papillae beginning on 
segment 8. Pygidium with anal ridge and 
pair of anal cirri. 

Hermilepidonotus robustus 

(Moore, 1905) 

Fig. 1 

Lepidonotus robustus Moore, 1905:544, pi. 
36:figs. 32-35.— Hartman, 1948:12. 

Lepidonotus helotypus. — Seidler, 1924:56 
(part). — Annenkova, 1937:145, pi. l:fig. 
1, pi. 3:fig. 14.— Hartman, 1938:109.— 
Uschakov, 1950:157; 1955:128, fig. 19, 
A-E; 1965:107, fig. 19, A-E.— Chlebov- 
itsch, 1961:165. Not Grube, 1877. 

Hermilepidonotus robustus. — Uschakov, 
1974:458, figs. A-H (synonymy); 1982: 
112, pi. 33, 1-8.— Buzhinskaja, 1985:82. 

Material examined. — Alaska: Shelikov 
Strait, 88-119 m, from hermit crab. Alba- 
tross sta 4291, 15 Aug 1903, holotype of 
L. robustus (USNM 5523). Virgin Bay, T. 
Kincaid, collector, 1 specimen (USNM 
3231 1). Coal Harbor, Unga Island, 1872, W. 
H. Dall, collector, 1 specimen (USNM 
18652). Kodiak Island, W. J. Fisher, collec- 
tor, 1 specimen (USNM 18867). Aleutian 
Islands, R. C. McGregor, collector, 1 spec- 
imen (USNM 18904). Canoe & Cold Bays, 
27-73 m, Sep & Oct 1940, 4 specimens 
(USNM 21318-21320; ZIASL, ident. by 
Hartman, 1948). 



Bering Sea: West Black Hill, 18 m, 27 
Jul 1957, Weber, collector, 1 specimen 
(USNM 32310). 

Okhotsk Sea: 16 Aug 1947, R V. Us- 
chakov, collector, 1 specimen (USNM 
43574, from ZIASL, as L. helotypus by Us- 
chakov, 1950). 

Description. — Length of holotype 37 
mim, width with setae 17 mm, 26 segments. 
Lengths of specimens examined 13^0 mm, 
widths 7-16 mm. Uschakov (1974) report- 
ed lengths to 65 mm, widths with setae to 
20 mm. Body elongate-oval, flattened dor- 
soventrally. Segments multiringed (wrin- 
kled), with pigmented transverse bands. 
Large oval elytra leaving middorsum un- 
covered, without fringes of papillae. Elytra 
rather thick, soft, of gelatinoid texture, with 
pigmented network on median and poste- 
rior areas, colorless on anterior and lateral 
areas; surfaces with slightly raised, bulbous, 
white spots with small "papillae" (thick- 
ened cuticle?) in center and scattered mi- 
cropapillae (Fig. IJ; Moore 1905, pi. 36:fig. 
32; Uschakov 1974, figs. C, D; 1982, pi. 
33,3). Dorsal cirri on non-elytrigerous par- 
apodia with cylindrical cirrophores, bulbous 
basally, on posterodorsal sides of notopo- 
dia, with styles long, cylindrical, with fila- 
mentous tips, and extending beyond neu- 
rosetae; dorsal tubercles forming slightly 
inflated areas (Fig. IG; Moore 1905, pi. 36: 
fig. 35; Uschakov 1974, fig. B; 1982, pi. 33, 
2). 

Bilobed prostomium with stout cerato- 
phore of median antenna in anterior notch, 
style rather short, with filamentous tip; lat- 
eral antennae with ceratophores formed of 
anterior extensions of prostomium, equal in 
length and lateral to ceratophore of median 
antenna, with styles slightly shorter; palps 
stout, tapering, slightly longer than anten- 
nae; anterior pair of eyes in region of great- 
est width of prostomium, larger than pos- 
terolateral pair; tentaculophores with few 
(0-7) slender setae on inner sides; dorsal 
and ventral tentacular cirri similar to me- 
dian antenna (Fig. lA; Uschakov 1974, fig. 
A; 1982, pi. 33, 1). 



VOLUME 109, NUMBER 1 



145 




Fig. 1. Hermilepidonotus robustus. A, D-J, holotype of Lepidonotus robustus (USNM 5523); B, C, pharynx 
from specimen from Aleutian Is. (USNM 18904): A, Dorsal view of anterior end, posterior pair of eyes partially 
hidden by segment II; style of left dorsal tentacular cirrus missing; setae lacking on right tentaculophore; B, 
Muscular pharynx removed from body and cut open midventrally and flattened, showing left and right ventral 
jaws on outside and dorsal pair of jaws in middle, and distal pharyngeal papillae; C, Pharyngeal branched 
papillae; D, Right elytrigerous parapodium from segment II, anterior view, acicula dotted; E, neuroseta from 
same; F, Right middle elytrigerous parapodium, anterior view, acicula dotted; G, Right middle cirrigerus para- 
podium, posterior view; H, Short and long notosetae from same; I, Short lower and longer upper neurosetae 
from same; J, Right middle elytron, with detail of micropapillae and pigmented network. Scales =1.0 mm for 
A, B; 0.2 nmi for C; 1.0 mm for D, F G; 0.1 mm for E, H, I; 1.0 mm for J. 



146 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Segment II without nuchal fold, with 
large bulbous elytrophores, biramous para- 
podia, and long ventral buccal cirri, lateral 
to ventral mouth, similar to tentacular cirri 
(Fig. ID; Uschakov 1974, fig. A; 1982, pi. 
33, 1). Notosetae similar to following seg- 
ments; neurosetae more slender than fol- 
lowing, with long spinous regions and long 
tapered bare tips (Fig. IE). Eversible mus- 
cular pharynx with 2 pairs of large hooked 
jaws and numerous border papillae enclos- 
ing numerous branched, threadlike papillae, 
forming thick brush (Fig. IB, C; Uschakov 
1974, figs. F, G, H; 1982, pi. 33, 6-8). 

Biramous parapodia with small subconi- 
cal notopodia on anterodorsal bases of larg- 
er neuropodia; smaller supraacicular part of 
neuropodia truncate and larger subacicular 
part diagonally truncate (Fig. IF, G; Moore 
1905, pi. 36:fig. 35; Uschakov 1975, fig. B; 
1982, pi. 33, 2). Notosetae forming close 
tuft, slender, with close transverse rows of 
fine teeth, some shorter, with blunt tips, and 
some longer, tapering to slender tips and ex- 
tending beyond tips of neuropodia (Fig. IF, 
H; Moore, 1905 pi. 36:figs. 34, 35). Neu- 
rosetae stout, dark golden color, forming 
dense bundles arranged in horizontal rows; 
2 supraacicular and 7 subacicular (Moore 
1905), with long spinous regions (13-20 
rows), supraacicular longer than subacicular 
ones, all with long, slightly hooked, bare 
tips (Fig. IF, I; Moore 1905, pi. 36:fig. 33; 
Uschakov 1974, figs. B, E, 1982, pi. 33, 2, 
5). Ventral cirri short, subulate, extending 
to lower tips of neuropodia (Fig. IF, G). 
Nephridial papillae beginning on segment 
8, on inflated areas and directed between 
parapodia. Pygidium with anus on middle 
of segments 25-26, with pair of anal cirri. 

Distribution. — Gulf of Alaska, Bering 
Sea, Aleutian and Kurile Islands, North Ja- 
pan and Okhotsk Seas, in 0-210 meters. 

Hermilepidonotus helotypus 

(Grube, 1877), new combination 

Fig. 2 

Polynoe (Lepidonotus) helotypus Grube, 
1877:49. 



Polynoe (Lepidonotus) gymnonotus Mar- 
enzeller, 1879:112, pi. l:fig. 3. 

Lepidonotus gymnonotus. — Mcintosh, 1885: 
64, pi. 10:fig. 4, pi. 17:fig. 5, pi. 9A:figs. 
2, 3. 

Lepidonotus holotypus. — Seidler, 1924:56, 
figs. 12, 13 (synonymy, part, not L. ro- 
bustus Moore). — Fauvel, 1933:8. — Mon- 
ro, 1934:358.— Not Annenkova, 1937: 
145 (=//. robustus)—i\2iVXmm\, 1938:109 
(part, not H. robustus). — Not Uschakov, 
1955:128 (=//. robustus). — Uschakov & 
Wu, 1959:10, 28-29; 1965:157; 1979:17 
(English translation). — Imajima & Hart- 
man, 1964:25 (synonymy). — Imajima & 
Gamo, 1970:2, fig. 4.— Rho & Song, 
1974:77, figs. 12-16; 1975:98.— Uchida, 
1982:3, figs. 4-7.— Jae, Lee, & Noh, 
1987:2, pi. lA-D. 

Lepidonotus {Lepidonotus) helotypus. — Us- 
chakov, 1982:106, pi. 30, 1-7). 

Material examined. — China: Chefdo, 
Grube, collector, syntype of P. (L.) heloty- 
pus (ZMB 1074). China, C. Ping, collector, 
2 specimens (BMNH 1925.5.5.11; USNM 
50910, ident. Monro, 1934). 

Japan: off Kobe, 15-91 m. Challenger 
Expedition, 2 Jul 1873, 3 specimens 
(BMNH 1885.12.1.46; 1921, 5,1,169, as L. 
gymnonotus by Mcintosh). Kobe, Gordon 
& Smith, collectors, 3 specimens (ZMB 
Q1858). Kokodate Bay, 2-25 m. Albatross 
sta 3656, 3657, 19 Sep 1895, 11 specimens 
(USNM 23749, 50911, 50912). Hokodate, 
Hilgendorf, collector, 2 specimens (ZMB 
2091, as P. (L.) gymnonotus). South Japan, 
N. Schedel, collector, 5 specimens (ZMH 
V-526, as L. gymnonotus by Augener). For- 
mosa or Japan, Laberer, collector, 5 speci- 
mens (ZMB 211, as L. gymnonotus). Hay- 
ama, Miura Peninsula, May 1966, Imajima, 
collector & identified, 1 specimen (USNM 
544770, from NSMT). 

Description. — Length of syntype (ZMB 
1074) 50 mm, width with setae 16 mm, seg- 
ments 26. Imajima & Hartman (1964) re- 
ported lengths to 63 mm, widths to 20 mm. 
Body elongate-oval, flattened dorsoventral- 



VOLUME 109, NUMBER 1 



147 




Fig. 2. Hermilepidonotus helotypus, specimen from Hokodate, Japan (USNM 50912); A, Dorsal view of 
anterior end, left lateral antenna small, regenerating; B, Right elytrigerous parapodium from segment II, antero- 
dorsal view, acicula dotted; C, Neuroseta from same; D, Right middle elytrigerous parapodium, anterior view, 
acicula dotted; E, Right middle cirrigerous parapodium, posterior view; F, short and long notosetae from same; 
G, Lower, middle and upper neurosetae from same; H, Right middle elytron, with detail of conical macropapillae 
and microtubercles among cellular network. Scales =1.0 mm for A; 0.5 mm for B, D, E; 0.1 mm for C, E G; 
1 .0 mm for H. 



ly. Segments multiringed, with transverse 
colored bands on posterior parts of seg- 
ments, matching elytra in color. Large oval 
elytra overlapping and leaving middorsum 



Intosh 1885, pi. 10:fig. 4). Elytra rather 
thick, soft, with pigmented network, except 
for anterior fourth and over scar of attach- 
ment to elytrophore; numerous microtuber- 



uncovered, without fringes of papillae (Mc- cles in cellular network of colorless area; 



148 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



surface of posterior pigmented area with 
projecting whitish conical macropapillae, 
variable in number and arrangement, from 
almost lacking to numerous (10-50, by Us- 
chakov 1982) (Fig. 2H; Mcintosh 1885, pi. 
17:fig. 5; Uchida 1982, fig. 5A, C-J; Us- 
chakov 1982, pi. 30, 2). Dorsal cirri on non- 
elytrigerous segments with cylindrical cir- 
rophores, bulbous basally, on posterodorsal 
sides of notopodia, with styles long, cylin- 
drical, with tapered tips and extending to 
tips of neurosetae; dorsal tubercles bulbous, 
inflated (Fig. 2E; Uchida 1982, fig. 6C). 

Bilobed prostomium with stout cerato- 
phore of median antenna in anterior notch, 
style with subterminal enlargement and fil- 
amentous tip; lateral antennae with cerato- 
phores formed of anterior extensions of 
prostomium, slightly shorter than ceratop- 
hore of median antenna; palps stout, taper- 
ing, longer than antennae; anterior pair of 
eyes in region of greatest width of prosto- 
mium, slightly larger than posterolateral 
pair; tentaculophores with aciculum and 
slender seta on inner side, and dorsal and 
ventral tentacular cirri, similar to median 
antenna (Fig. 2A; Uchida 1982, figs. 5A, B, 
6A; Uschakov 1982, pi. 30, 1). 

Segment II with slightly developed nu- 
chal fold, with large bulbous elytrophores, 
biramous parapodia, and long ventral buc- 
cal cirri, similar to tentacular cirri (Fig. 2A, 
B; Uchida 1982, fig. 5B). Notosetae similar 
to following segments; neurosetae more 
slender than following, with long spinous 
regions and slender entire tips (Fig. 2C). 
Eversible pharynx with 13-17 pairs of mar- 
ginal papillae (Uschakov 1982, pi. 30, 7). 

Biramous parapodia with small conical 
notopodia on anterodorsal sides of large 
neuropodia; neuropodia with subequal pre- 
setal and postsetal lobes; smaller supraaci- 
cular part of neuropodia truncate and larger 
subacicular part diagonally truncate (Fig. 
2D, E; Uchida 1982, fig. 6B, C). Notosetae 
forming small bundle, thin, with long spi- 
nous regions, some short, with blunt tips 
and some long, tapering to fine tips (Fig. 
2F; Uchida 1982, fig. 7E, F). Neurosetae 



numerous, stout, dark reddish amber-col- 
ored, with long spinous regions, supraaci- 
cular ones with longer spinous regions than 
subacicular ones, all with rather long, 
slightly hooked, entire tips (Figure 2G; 
Uchida 1982, fig. 7I-K). Ventral cirri short, 
subulate, not extending to lower tips of neu- 
ropodia (Fig. 2D, E; Uschakov 1982, pi. 30, 
4). Nephridial papillae beginning on seg- 
ment 8. Pygidium with anus medial to par- 
apodia of segment 25, with pair of anal cirri 
medial to dorsal cirri of last 2 segments, 
directed posteriorly. 

Distribution. — Yellow Sea, China, South 
Japan, Korea, Kurile Islands, in 0-108 me- 
ters. 

Remarks. — Uschakov referred Lepidon- 
otus robustus to the new genus Hermilepi- 
donotus, based mainly on the unusual struc- 
ture of the pharynx, with numerous thread- 
like papillae forming a thick brush. Speci- 
mens of Lepidonotus, and the majority of 
other polynoids, have the pharynx with 9 
pairs of large marginal papillae. Lepidono- 
tus helotypus, referred herein to Hermile- 
pidonotus, has more numerous border pa- 
pillae (13-17 pairs) than the 9 pairs in other 
species of Lepidonotus. Also, the neuropo- 
dia of the two species of Hermilepidonotus 
differ from those of other Lepidonotus spe- 
cies in having subequal presetal and postse- 
tal lobes (Figs. IF, G, 2D, E) and lacking 
the larger presetal conical acicular process 
as found in species of Lepidonotus. 

Literature Cited 

Annenkova, N. P. 1937. [Polychaeta Fauna of the 
Northern Part of the Japan Sea]. — Leningrad 
Gosudarstrennyi Gidrologicheskii Institut Issle- 
devaniia moorei SSSR, 23:129-215. [In Rus- 
sian, English summary.] 

Buzhinskaja, G. N. 1985. [Polychaete worms (Poly- 
chaeta) of the shelf off south Sakhalin and their 
ecology.]. — Issledovaniia Fauny morei Zoolo- 
gischeskii Institut Academiia Nauk SSSR 
30(38):72-224. [In Russian.] 

Chelbovitsch, V. V. 1961. [Polychaete worms (Poly- 
chaeta) of the littoral Kurile Islands.]. — Aka- 
demiia Nauk SSSR, Issledovaniia Dalvesvos- 
tochnykh Morei SSSR, 7:151-260. [In Rus- 
sian.] 



VOLUME 109, NUMBER 1 



149 



Fauvel, P. 1933. Annelides Polychetes du Golfe du 
Pel Tcheu Ly. — Publication Musee Hoangho 
Paiho de Tien-Tsin, No. 15:1-67. 

Grube, H. 1877. Ueber einer Sammlung von wirbel- 
losen Seethieren, welche Herr Dr. Eugen Rei- 
mann dem hiesigen zoologischen Museum zum 
Geschenk gemacht. — Jahres-Bericht der Schles- 
ischen Gessellschaft fiir vaterlandische Cultur 
(1876) 54:48-51. 

Hartman, O. 1938. The types of the polychaete worms 
of the families Polynoidae and Polyodontidae in 
the United States National Museum and the de- 
scription of a new genus. — Proceedings of the 
United States National Museum 86(3046): 107- 
134. 

. 1948. The polychaetous annelids of Alas- 
ka. — Pacific Science, 2:3—58. 

Imajima, M., & S. Gamo. 1970. Polychaetous Anne- 
lids from the intertidal zone of Manazuru, Kan- 
agawa Prefecture. — Science Reports of the Yo- 
kohama National University, Section II(16):1- 
18. 

, & O. Hartman. 1964. The polychaetous an- 
nelids of Japan, Part I. — Allan Hancock Foun- 
dation Publications Occasional Paper, No. 26: 
1-257. 

Jae, J.-C, J.-H. Lee, & Y.-T. Noh. 1987. Taxonomic 
study on polynoid polychaetes in Korea. 2. Sub- 
family Lepidonotidae. — Bulletin of the Korean 
Fisheries Society 20(1): 1-1 5. [In Korean, En- 
glish summary.] 

Kinberg, J. G. H. 1856. Nye slagten och arter af An- 
nelider — Ofversigt af Konglia Vetenskaps-Aka- 
demiens Forhandlingar, Stockholm 12:381-388. 

Marenzeller, E. von. 1879. Siidjapanische Anneli- 
den. — Denkschriften der Kaiserlichen Akade- 
mie der Wissenschaften, Wien 41(2): 109-154. 

Mcintosh, W. C. 1885. Annelida Polychaeta. — In Re- 
port on the Scientific Results of the Voyage of 
the H.M.S. Challenger . . . 1873-76 . . . Zool- 
ogy 12(34): 1^42. 

Monro, C. C. A. 1934. On a collection of Polychaeta 
from the coast of China. — Annals and Maga- 
zine of Natural History, series 10, 13:353-380. 

Moore, J. P. 1905. New species of Polychaeta from 
the North Pacific, chiefly from Alaskan wa- 
tes. — Proceedings of the Academy of Natural 
Science, Philadelphia, 57:525-554. 

Rho, B. J., & K. H. Song. 1974. A Study on the Clas- 
sification of the Korean Polychaeta. — Journal of 



Korean Research Institute for Better Living, 12: 
73-85. 
-. 1975. On the Classification and Distribution 



of the Marine Benthic Animals in Korea. 2. 
Polychaetous annelids. — Journal of Korean Re- 
search Institute for Better Living 14:95-118. 

Seidler, H. J. 1924. Beitrage zur Kenntnis der Poly- 
noiden, I. — Archiv fiir Naturgeschichte 89A, 
Heft 11:1-217. 

Uchida, H. 1982. Polychaetous annelids from Kishu. 
5. Aphroditoidea (4). — Nanki Seibutsu 24(1):1- 
7. [In Japanese, English summary.] 

Uschakov, P. V. 1950. [Polychaete worms (Polychae- 
ta) from the Okhotsk Sea.]. — Issledovaniia 
Morei dalnevost Morei SSSR, 2:140-234. [In 
Russian, English summary.] 

. 1955. [Polychaeta of the Far Eastern Sea of 

the U.S.S.R. In Keys to the Fauna of the 
U.S.S.R.]. — Zoological Institute of the Acade- 
my of Sciences of the U.S.S.R. No. 56:1^45. 
[In Russian; 1965, English translation by the Is- 
rael Program for Scientific Translations, 419 

PP] 

. 1974. [The aberrant type of pharyngeal pa- 
pillae in polynoids {Hermilepidonotus gen. n., 
Polynoidae, Polychaeta).]. — Zoologischesky 
Zhurnal, 53(3):457^60. [In Russian, English 
summary.] 

. 1982. [Polychaetes of the suborder Aphrod- 

itiformia of the Arctic Ocean and the North- 
western part of the Pacific, Families Aphrodi- 
tidae and Polynoidae.] In Fauna of the USSR 
Polychaetes, Vol. 2, Part 1. Academy of Sci- 
ences of the USSR Zoological Institute, 271 
pages. [In Russian; English translation by Na- 
tional Museum of Canada, not published.] 

, & B. L. Wu. 1959. The Polychaetous An- 
nelids of the Families Phyllodocidae and 
Aphroditidae from the Yellow Sea. — Archiv In- 
stituta Oceanologia Sinica 1(4): 1^0. 

, & . 1965. [Polychaeta Errantia of the 



Yellow Sea. Akademia Nauk SSSR, Zoological 
Institute.] In Fauna of Seas of Northwestern part 
of Pacific Ocean. — Isslededovaniya Fauny Mor- 
ei 3(11): 145-258. 1979. English translation, 
U.S. Depart. Comm. Nat. Tech. Inform. Serv. 
Springfield, Virginia, 137 pages. 
Willey, A. 1902. Polychaeta. In Report on the collec- 
tions of natural history made in the Antarctic 
regions during the voyage of the Southern 
Cross, London 12:262-283. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(1): 150-203. 1996 

The Antarctic genus Callozostron and its relationship to Primnoella 
(Octocorallia: Gorgonacea: Primnoidae) 

Frederick M. Bayer 

Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — Specimens of the gorgonacean genus Callozostron obtained in 
Antarctic waters during operations conducted by the U.S. Antarctic Research 
Program reveal that Callozostron horridum Ktikenthal, 1909, is a junior syn- 
onym of Callozostron mirabile Wright, 1885. Callozostron carlottae Ktiken- 
thal, 1909, is confirmed as a valid species, and two new species, Callozostron 
diplodiadema and Callozostron acanthodes are described. All species are il- 
lustrated by stereoscopic scanning electron micrographs (SEM). The genus 
Primnoella Gray, 1858, is restricted to the "Compressae" species group, and 
polyps and sclerites of Primnoella australasiae (Gray, 1850), type species of 
the genus, are illustrated by SEM for the first time. A new genus, Convexella, 
is established for the "Convexae" species group of Primnoella, and polyps of 
Primnoella magelhaenica Studer, 1879, type species of Convexella, from a 
wide bathymetric and geographic range are illustrated by SEM to show mor- 
phological variation within the species. 



During biological operations of the U.S. 
Antarctic Research Program (USARP), sev- 
eral deep-water dredge and trawl tows in 
Antarctic waters have yielded specimens of 
the three species of Callozostron so far de- 
scribed, C. mirabile Wright, 1885, C. car- 
lottae Ktikenthal, 1909, and C horridum 
Ktikenthal, 1909. This material shows that 
C. horridum cannot be maintained as a spe- 
cies separate from C mirabile, and that a 
species distinct from C. carlottae remains 
to be described, in addition to a new species 
having a branched colony unlike any 
known heretofore. These are here described 
respectively as Callozostron diplodiadema 
and C acanthodes, new species. 

Operations of USARP in the Patagonian 
region and on Burdwood Bank have ob- 
tained numerous colonies attributable to 
Primnoella magelhaenica Studer as report- 
ed by Thomson & Ritchie (1906, as ma- 
gellanica) from the Scotia cruise and by 
Broch (1965) from the Brategg expedition. 
These show sufficient variation to accom- 



modate both Primnoella magelhaenica and 
P. flagellum as described by Studer (1879) 
from the same geographical area. The larg- 
est colonies are superficially almost indis- 
tinguishable from specimens of Callozos- 
tron diplodiadema new species and C car- 
lottae Ktikenthal, obscuring but not elimi- 
nating the distinction between Primnoella 
and Callozostron. 

Preparation of samples. — In general, 
sclerites, intact whorls and isolated polyps 
were prepared according to methods de- 
scribed elsewhere (Bayer & Stefani 1989: 
450-451). However, in order to determine 
the arrangement of opercular and circumo- 
percular sclerites, single polyps were indi- 
vidually macerated in Sodium hypochlorite 
solution and observed under the dissecting 
microscope during the process in order to 
distinguish opercular from circumopercular 
scales, and the isolated sclerites washed one 
by one by tranferring them through a series 
of drops of distilled water, drying, and 
mounting in sequence on SEM stubs. 



VOLUME 109, NUMBER 1 



151 



Callozostron P. Wright, 1885 

Callozostron P. Wright, 1885:691.^Studer 
[& Wright], 1887:48.— Wright & Studer, 
1889:48.— Versluys, 1906:124.— Kiiken- 
thal, 1912:331; 1915:152; 1919:449; 
1924:306.— Bayer, 1981:936 (in key 
only).— Bayer & Stefani, 1989:455 (in 
key only). (Type species: C. mirabile 
Wright & Studer, 1889, by subsequent 
monotypy.) 

Diagnosis. — Flagelliform, unbranched or 
sparsely branched primnoids with tall, cy- 
lindrical polyps standing almost vertically 
from the axis, arranged in whorls; bases of 
polyps may be partially fused to form un- 
branched polyp leaves; opercular scales of 
nearly uniform size, distinctly differentiated 
from marginal scales; at least 4 marginal 
scales with very long, slender apical spine; 
body completely covered by scales. No ir- 
regular, tuberculate sclerites in walls of lon- 
gitudinal stem canals. 

Remarks.- — In my key to octocoral gen- 
era (Bayer 1981:936), Callozostron is not 
distinguished from Ainigmaptilon, in both 
of which the polyps partially fuse to form 
pennatulacean-like "polyp-leaves." How- 
ever, the spiculation of the polyps differs in 
the two genera. All nominal species of Cal- 
lozostron have 8 well-differentiated smaller 
opercular scales within a ring of large mar- 
ginal scales bearing long apical spines. In 
Ainigmaptilon the distalmost 8 scales are 
larger than the marginals, which are not 
well differentiated from the body scales be- 
low. Those of A. antarcticum (Molander) 
are triangular, with a broadly pointed apex 
(Molander 1929: fig. 22a) reminiscent of 
the opercular scales of Primnoella gracilis 
Molander (1929: fig. 18a), and those of A. 
virgularoides (Molander) are lancet-shaped 
with a short apical spine. In A. edisto Bayer 
(1950: fig. 2a) they bear long, smooth 
spines, and in A. haswelli Dean they have 
a strong, serrated apical spine (Thomson & 
Rennet, 1931: pi. 13, fig. 1). The 8 distal- 
most polyp scales of A. antarcticum and A. 
virgularoides converge over the infolded 



tentacles in a manner similar to those of 
Primnoella gracilis Molander, but in A. 
haswelli and A. edisto these scales cannot 
fold over the tentacles because of their long 
apical spines. Molander (1929), Carlgren 
(1943) and Bayer (1950) call these sclerites 
"Deckschuppen" or "opercular scales" 
whether or not they have long spines but, 
in fact, there is no functional "operculum" 
as in Callozostron. 

In the examples so far reported, the axis 
of Ainigmaptilon terminates basally in a 
funnel-shaped expansion which encloses a 
quantity of mud that served to anchor the 
colony in soft substrate, whereas in those 
cases that are known, the axis of Callozos- 
tron terminates in a small, discoidal hold- 
fast for attachment to solid substrate. 

Most records of Ainigmaptilon are from 
shallow or moderate depths: wallini: 75 m; 
antarcticum, 125 m; haswelli: 517—549 m; 
edisto: 182 m; USNM records of antarcti- 
cum: 88-526 m; edisto: 182-662 m. 

Except for one doubtful record, speci- 
mens of Callozostron have been taken at 
much greater depths, usually below 1000 m 
and mostly between 2000 and 3000 m. 

Accordingly, there seems to be no doubt 
that both Callozostron and Ainigmaptilon 
should be treated as valid genera separable 
on the basis of differences in the opercular 
and circumopercular scales as mentioned 
above. 

In the discussion of his new genus Ly- 
curus (— Ainigmaptilon), Molander (1929: 
70) pointed out the similarities of the genus 
to Primnoella gracilis and suggested that it 
may have arisen from such a form. A strong 
morphological similarity also exists be- 
tween Callozostron and certain Primnoella 
species including P. magelhaenica Studer 
and P. flagellum Studer. 

In Callozostron carlottae Kiikenthal and 
C. diplodiadema new species, the basal fu- 
sion of adjacent polyps is minimal (see 
Figs. 9, 11, and KUkenthal 1912:334, fig. 
43), inviting comparison with Primnoella 
magelhaenica Studer, in which the polyps 
are not appressed to the axis and often (but 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



not always) stand out almost vertically (see 
Figs. 21, 23, 27, and Broch 1965: pi. 1, fig. 
1). Primnoella magelhaenica has distinctly 
pointed marginal scales that surround and 
overlap the opercular scales proper. In this 
respect, the polyps of P. magelhaenica are 
similar to those of C. carlottae, differing 
most conspicuously in the absence of 
strongly developed apical spines on the 
scales of three whorls surrounding the oper- 
culum, which is poorly differentiated from, 
and partially covered by, the marginal (cir- 
cumopercular) sclerites. 

Key to species of Callozostron 

1(2). Colonies dichotomously branched, 
polyps arranged in whorls. Whorls 
composed of up to 6 polyps. Apex of 
opercular scales prolonged as a nar- 
row spine. Marginal spines smooth, 
about 7X as long as height of basal 
part acanthodes, new species 

2(1). Colonies flagelliform, sometimes bi- 
furcate but not repeatedly branched, 
polyps arranged in whorls that may be 
densely crowded. Apex of opercular 
scales more or less acutely pointed 
but not prolonged as a narrow spine. 

3(4). Whorls sometimes densely crowded 
and more or less obscured, composed 
of as many as 12 or even more pol- 
yps; polyps about 3 mm tall exclusive 
of marginal spines; marginal spines 
smooth, slender, 4-6, commonly 4, 
developed only on the marginal trans- 
verse row of body scales, 2-3 X as 
long as the height of the wide basal 
part mirabile 

4(5). Colonies flagelliform or bifurcate. 
Whorls distinct, composed of 8-9 
polyps; polyps about 2 mm tall exclu- 
sive of marginal spines; marginal 
spines stout, with raised longitudinal 
ridges, about 24, developed on 3 
transverse rows of body scales, about 
2X as long as height of basal part 
carlottae 

5(1). Colonies flagelliform. Whorls dis- 
tinct, composed of up to 14 polyps; 
polyps about 2 mm tall exclusive of 
marginal spines; marginal spines slen- 



der, smooth, about 16, developed on 
2 transverse rows of distal body 
scales, about 5X as long as height of 
basal part . . .diplodiadema, new species 

Callozostron mirabile Wright, 1885 
Figs. 1, left; 2-6 

Callozostron mirabile Wright, 1885:691, 
figs. 234, 235.— Wright & Studer, 1889: 
48, pi. 10, figs. 1-5; pi. 20, fig. 1.— Ver- 
sluys, 1906:124.— Kukenthal, 1912:328- 
333 passim; 1919:450; 1924:306.— 
Schimbke, 1915:22. 

Callozostron horridum Kukenthal, 1909:49 
1912:331, figs. 38^2, pi. 22, figs. 12, 13 
1919:451; 1924:307.— Schimbke, 1915 
23. 

Material. — Antarctica, Oates Coast: 
64°59'S, 160°36'E, 2836-2864 m, USNS 
Eltanin sta. 1957, 7 Feb 1967; one colony 
USNM 78662. South Atlantic Ocean, be- 
tween South Orkney Islands and Palmer 
Peninsula: 63°03'S, 49°11'W, 2653-2941 
m, USNS Eltanin sta. 529, 3 Mar 1963; 2+ 
colonies USNM 77377 (SEM 1159). South 
Atlantic Ocean, off South Georgia Island: 
56°04'S, 33°59'W, 3239-3138 m, USNS El- 
tanin sta. 722, 8 Sep 1963, 5+ colonies 
USNM 77378 (SEM 1160, 1166, 1167, 
1622, 1623). South Atlantic Ocean, off 
South Georgia Island: 54°00'S, 33°40'W, 
2718-2663 m, USNS Eltanin sta. 723, 9 
Sep 1963. 1 colony USNM 77381. South 
Atlantic Ocean, off South Sandwich Is- 
lands: 57°22'36"S, 26°34'00"W, 2248-2402 
m, RA^ Islas Orcadas sta. 51, 26 May 
1975, 1+ colony USNM 82876 (SEM 
1624, 1625). South Atlantic Ocean, off 
South Sandwich Islands: 57°39'24"S, 
26°26'42"W, 415-612 m, RA^ Islas Orca- 
das sta. 52, 26 May 1975, 1+ colonies 
USNM 77339 (probably a contaminant 
from sta. 51). South Pacific Ocean, off An- 
tipodes Island, New Zealand: 49°40'S, 
178°56'E, 952-1336 m, USNS Eltanin sta. 
1852, 3 Jan 1967, 3+ colonies USNM 
77402. Scotia Sea: 60°02'S, 49°14'W, 
3819-3876 m, USNS Eltanin sta. 545, 7 



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153 




Fig. 1. Left, Callozostron mirabile Wright, USNM 82876. Colony with axis still in place but broken; Right, 
Callozostron diplodiadema new species, USNM 77386. Two incomplete colonies (syntypes). 



Mar 1963, 1 incomplete colony ripped from 
axis and broken, USNM 85294 (SEM 
1630-1634). 

Diagnosis. — Callozostron with up to 12 
or more polyps in whorls that may be ob- 
scured by close crowding. Long, slender 
apical spine developed on 4-6 marginal 
scales of polyps. 

Remarks. — The distalmost whorl in a 
colony may consist of as few as 4 polyps 
(Fig. 3), but this number quickly increases 
proximad. Fully developed polyps (Fig. 2, 
top) are about 3 mm tall excluding the 4-6 
long, marginal spines (Fig. 2, bottom). 



which usually are more or less broken, and 
indistinctly united by their bases into 
groups of a few individuals (Fig. 2, top). In 
contraction, eight distalmost polyp scales 
fold over the retracted tentacles as a conical 
operculum (Fig. 4). The marginal scales of 
immature polyps (Fig. 5) do not have the 
long apical spine present on fully developed 
individuals (Figs. 2, 4). Sclerites (Fig. 6) 
are of the generalized primnoid type: thin, 
rounded scales with finely dentate margins, 
smooth externally and covered with closely 
set complex tubercles internally, those of 
the polyp body (Fig. 6c) larger than those 



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Fig. 2. Callozostron mirabile Wright, USNM 85294. Top, Four polyps from a whorl; Bottom, Detail of oral 
end of polyp with four marginal scales prolonged into spines. Stereoscopic pairs (SEM 1631). 



of the coenenchyme (Fig. 6d); the opercular 
scales are of the usual roughly triangular 
shape, thin, with a broad, shallow apical 
groove. Four to six of the marginal scales 



are unusual in having an exceptionally long, 
thin, fragile, apical spine (Fig. 6b). 

The present material provides evidence 
that the coenenchyme and polyps of the 



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155 




Fig. 3. Callozostron mirabile Wright, USNM 85294; whorl with only four polyps, from distal end of colony. 
Top, Oral view; Bottom, Side view. Stereoscopic pairs (SEM 1632). 



original type specimen may have been 
stripped from the axis during collection and 
the whorls tightly compressed together as 
shown in the original figures. Some of the 
specimens here reported are similarly dam- 



aged. Some have a well-developed axis 
(Fig. 1), and some have been stripped off 
of the axial skeleton during collection. Fur- 
thermore, as specimens of Callozostron 
carlottae Kiikenthal, Primnoella magel- 



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Fig. 4. Callozostrori mirabile Wright, USNM 85294. Top, Oral view of polyp showing opercular scales in 
closed position; Bottom, Oblique view of polyp with opercular scales in closed position. Stereoscopic pairs 
(SEM 1632). 



haenica Studer, and Ophidiogorgia para- 
doxa Bayer have been similarly stripped 
from their supporting axes, this circum- 
stance seems not to be rare among unbran- 
ched primnoids. 



Callozostrori mirabile appears to be 
widely distributed around Antarctica. The 
type locality is off the Leopold & Astrid 
Coast of Princess Elizabeth Land at 
65°42'00"S, 79°49'00"E in 3036 m. It also 



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157 




Fig. 5. Callozostron mirabile Wright, USNM 85294. Lateral and oral views of young polyp before differ- 
entiation of marginal and opercular scales. Stereoscopic pairs (SEM 1633). 



has been taken by USNS Eltanin in the vi- 
cinity of the Balleny Islands off Wilkes 
Land, 64°59'S, 160°36'E, in 2536-2864 m; 
off the Antipodes Is., New Zealand, at 
49°40'S, 178°56'E, in 952-1336 m; by R/ 
V Islas Orcadas at 2 stations in the Scotia 



Sea at 2653-2941 m and 3819-3876 m, and 
2 stations off South Sandwich Is. at 224— 
2402 m and 415^62 m. The unusually 
shallow depth of the specimen from sta. 52 
(USNM 77339) is suspect, because it im- 
mediately followed sta. 5 1 of the same date 



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159 



in 2248-2402 m (USNM 82876) and prob- 
ably was obtained with the same Blake 
trawl. The probability of contamination in 
sta. 52 of fragments from sta. 51 must be 
considered sufficiently strong to discount 
the anomalous bathymetric record. 

Callozostron carlottae Kiikenthal, 1909 
Figs. 7, top; 8-10 

Callozostron carlottae Kiikenthal, 1909:49; 
1912:334, figs. 43-47, 53, 54, pi. 22, 
figs. 14-17; 1919:451; 1924:307, fig. 
168.— Schimbke, 1914:25. 

Material. — Antarctic Ocean, off South 
Orkney Islands: 61°04'S, 39°55'W, 2355- 
2897 m, USNS Eltanin sta. 1545, 11 Feb 
1966. 2 bifurcate colonies lacking holdfast, 
and 1 simple branch, USNM 77387. (SEM 
1626, 1627.) 

Diagnosis. — Flagelliform or bifurcate 
Callozostron with polyps in whorls of 8 or 
9, having 2 rings of 8 scales (marginal and 
submarginal) surrounding the operculum, 
most or all with a stout apical spine; basal 
fusion of polyps negligible. 

Comparisons. — This species superficial- 
ly resembles Callozostron diplodiadema 
new species, but the polyps are shorter and 
the spines of the marginal and submarginal 
scales are shorter, stouter, distinctly tapered 
and prismatic in section. 

Remarks. — The specimens here recorded 
agree with the type material of C carlottae 
insofar as the ambiguities of the original de- 
scription allow, except in growth form. The 
type specimen (Kiikenthal 1912) was fla- 
gelliform and showed no evidence of 
branching, but two of the three colonies in 
the present collection are bifurcate imme- 
diately above the holdfast (Fig. 7, top) and 
the third is a single flagelliform branch that 
could be half of a bifurcate colony. Super- 
ficially, an unbranched colony would close- 



ly resemble a stout colony of Primnoella 
magelhaenica Studer, 1879 (cf. Figs. 20, 
21). 

Polyps are arranged in regular whorls 
(Fig. 8, top) of 9 (in the original type ma- 
terial mostly 8, but 9 in the median whorls, 
an insignificant difference); their overall 
height in the present material (Fig. 9) is 
about 2.5 mm (2.3-2.7 mm), of which a 
little more than 0.5 mm consists of the pro- 
jecting marginal spines, compared with 1.2 
mm overall as given by Kiikenthal (1912: 
335). The discrepancy may be explained by 
a difference in the way measurements were 
made but this cannot be verified as neither 
magnification nor scale accompanies Kii- 
kenthal's illustrations (1912: figs. 43, 53). 
In shape, the polyps (Fig. 9) are strikingly 
similar to Kiikenthal's drawing (1912: fig. 
43), with a double ring of strong spines sur- 
rounding the operculum (Fig. 8, bottom). 
Basal fusion of polyps is no more than oc- 
curs in Primnoella magelhaenica (cf. Fig. 
24). 

The sclerites are of the usual primnoid 
form, but somewhat thicker than in Callo- 
zostron mirabile. The body scales are 
broadly oval (Fig. 10c), and the opercular 
scales (Fig. 10a) diminish in size toward the 
adaxial side more than is the case in mira- 
bile. The apical spine of the marginal and 
submarginal scales is marked by a few 
smooth longitudinal ridges (Fig. 10b). Al- 
though Kiikenthal stated that numerous 
warts cover the surface of the lower part of 
the marginal scales, it is not clear whether 
he referred to the outer surface, the inner 
surface, or both. In primnoids, the inner 
surface of sclerites is invariably covered 
with complex warts, but the outer surface 
varies according to species and/or location. 
His illustration of a marginal scale (1912: 
fig. 44) appears to show the inner surface, 
leaving the nature of the outer surface un- 



Fig. 6. Callozostron mirabile Wright, USNM 85294. Sclerites (SEM 1634). a. Opercular scales; b. Marginal 
scales; c. Body scales; d, Coenenchymal scales. 



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161 



Specified. In the present material, the outer 
surface of the proximal part of both mar- 
ginal and opercular scales is covered with 
simple granules (Fig. 10a, b) while that of 
the body scales is smooth except for the 
proximal edge (Fig. 10c). 

Callozostron diplodiadema, 

new species 

Figs. 1, right; 11-14 

Material. — Scotia Sea, SW of South 
Georgia Island: 55°01'S, 39°55'W, 2886- 
3040 m, USNS Eltanin sta. 1537, 8 Feb 
1966. 7+ colonies USNM 77386. (SEM 
1628, 1655-57.) 

Diagnosis. — Flagelliform Callozostron 
with polyps in distinct whorls of up to 14, 
having 2 rings of 8 scales (marginal and 
submarginal) surrounding the operculum, 
each with a long, smooth apical spine; basal 
fusion of polyps negligible. 

Description. — The material consists of 7 
incomplete colonies still with supporting 
axial skeleton, the longest of which is 19 
cm in length lacking holdfast (Fig. 1, right), 
and 10 pieces of various size stripped off 
the axis during collection. It is likely that 
at least some of these pieces occupied na- 
ked spaces on some of the damaged colo- 
nies. One colony with axis nearly or quite 
intact apically is 9.7 cm long and attached 
by a small discoidal holdfast to a flat black 
pebble. The polyps stand almost perpendic- 
ular to the axis and are arranged in regular 
whorls (Figs. 11, 12), mostly of 9-12 but 
initially of 4 and ultimately as many as 14. 
Where the coenenchyme remains undis- 
turbed on the axis, 4-5 whorls occur in 1 
cm of length, but where artificially com- 
pressed along the axis or completely 
stripped from it, 7 whorls may be closely 
crowded in 1 cm of length. 

The polyps are about 2 mm tall exclusive 
of the operculum and marginal spines (Fig. 



12, bottom), only the basal part of their bod- 
ies fused together, the free distal half about 
0.5 mm in diameter widening distally to 
about 0.8 mm at the level of the circumo- 
perculum. The marginal (circumopercular) 
and submarginal scales each have a long, 
smooth, somewhat curved apical spine (Fig. 
14b), together forming crown of 16 spines 
surrounding the operculum (Fig. 13, top). 
The operculum (Fig. 13, bottom) is com- 
posed of 8 tall, triangular scales of about 
equal size, with somewhat convex sides 
(Fig. 14a). The opercular scales are vertical- 
ly aligned with the circumopercular and fol- 
lowing longitudinal rows of body scales. 
The distal margin of some of the scales in 
the transverse row beneath the submarginals 
may be more or less acutely pointed but not 
produced as a spine (Fig. 14c). The abaxial 
and adaxial longitudinal rows of body scales 
consist of 5-6 scales each, including the spi- 
nous submarginals and circumoperculars. 
Their outer surface is smooth and glossy, the 
inner surface covered by complex tubercles 
where embedded in the body wall (Fig. 
14d). The coenenchymal sclerites (Fig. 14e) 
are rounded scales smaller than those of the 
polyps. The tentacles are devoid of sclerites. 
No irregularly tuberculate sclerites are pres- 
ent in the walls separating the longitudinal 
stem canals. 

Comparisons. — This species superficial- 
ly resembles C. carlottae but the polyps are 
taller and the spines of the marginal and 
submarginal scales are much longer, 
smooth, cylindrical or elliptical in cross 
section. 

Etymology. — Greek SlttXoos, double + 
8Ld8T||xa, diadem, crown. Noun in apposi- 
tion. 

Callozostron acanthodes, new species 
Fig. 7, bottom; 15-17 

Material.— East of North Island, New 
Zealand, 38°24'S, 178°53'E, 1354-1995 m. 



Fig. 7. Top, Callozostron carlottae Kiikenthal, two bifurcate colonies, USNM 77387; Bottom, Callozostron 
acanthodes, holotype, USNM 94575. 



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162709 5.0KV X40.0 .75mm 



162711 5.0KV X40.0 .75mm 



Fig. 8. Callozostron carlottae Kiikenthal, USNM 113^1. Top, Part of branch showing two whorls of polyps 
(SEM 1626); Bottom, Oral view of polyp (SEM 1627). Stereoscopic pairs. 



USNS sta 1712, 5-foot Blake trawl 40 min- Diagnosis. — Dichotomously branched 

utes at depth, 28 May 1966. One colony Callozostron with tall, slender polyps ar- 

with branches detached, USNM 94575. ranged in widely spaced whorls commonly 

(SEM 1594, 1595, 1603, 1617, 1618.) of 5, sometimes 6, rarely isolated; usually 



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163 



4 whorls in 1 cm; individuals often partly 
united basally as pairs. Marginal scales with 
extremely long, slender spine, vertically 
aligned with opercular scales; apex of oper- 
cular scales prolonged as a slender, acute 
spine. 

Description. — The holotype colony (Fig. 
7, bottom) is sparsely dichotomously 
branched in one plane, 1 1 cm tall lacking 
the holdfast and perhaps the uppermost ter- 
minal twig; undivided terminal twigs to 5 
cm in length. The axis is calcified, light 
brown with little or no luster in the main 
stem, cylindrical, smooth, 0.9 mm in di- 
ameter, becoming paler distad; in the ter- 
minal branches it is cream white with mod- 
erate pearly luster, proximally about 0.5 
mm in greater diameter, flattened in the 
plane of branching, tapering distad to an ex- 
tremely fine tip. 

The polyps are situated in widely spaced 
whorls of 3-6, commonly 5 (Fig. 15); usu- 
ally 4 whorls in 1 cm except at the tips of 
branches where there may be 5. The distal- 
most whorl consists of 2-4 polyps, new 
polyps arising within whorls proximad. No- 
where in the colony is there any evidence 
of new whorls of polyps originating be- 
tween fully developed whorls, indicating 
that growth is exclusively terminal. Rarely 
a single polyp occurs in the location of a 
whorl. 

Fully developed polyps (Fig. 15, bottom) 
are about 2.5 mm tall and 0.5 mm in di- 
ameter, nearly cylindrical, somewhat flared 
immediately below the tentacles, and in- 
clined slightly distad. Adjacent polyps may 
be proximally somewhat fused to form 
pairs (Fig. 16, top). They are covered by 
roughly fanshaped scales aligned in more or 
less distinct longitudinal rows consisting of 
about 10 scales. The body scales (Fig. 17a) 
are externally smooth, internally sculptured 
with scattered complex tubercles; their dis- 
tal free margins are irregularly serrate. Sev- 
eral of the marginal scales (Fig. 17b) have 
an extremely long, slender apical spine that 
may exceed 3 mm in length; the spine usu- 
ally is smooth, but in some cases shows 



traces of widely scattered, low, simple gran- 
ules. Owing to their extreme delicacy, none 
of the marginal scales were preserved with 
spine intact, so their greatest length cannot 
be determined. The operculum (Fig. 16, 
bottom) consists of eight ovate scales with 
the distal margin smooth and apically ex- 
tended as a smooth, slender spine (Fig. 
17c). 

The coenenchyme is thin, covered by 
small, externally smooth, rounded scales 
(Fig. 17d). 

Comparisons. — Callozostron acanthodes 
new species differs sharply from previously 
known species by its openly dichotomous 
branching, the wide spacing of the whorls, 
and the conspicuous apical spine of the of 
the opercular scales. 

Etymology. — Greek aKavScoStis, thorny. 

Remarks. — This species is assigned to 
the genus Callozostron because of the reg- 
ular verticillate arrangement of the polyps, 
clear differentiation of opercular from mar- 
ginal scales, spinelike apical projection of 
the marginal scales, and regular rows of 
body scales. The branched colonial form is 
not considered an exclusive character as it 
is here shown that colonies of C carlottae 
may be bifurcate. The polyps bear some 
resemblance to those of Mirostenella Bay- 
er, 1988, but are proportionally much 
taller, with several but not all of the mar- 
ginal scales furnished with a strong apical 
spine. 

Discussion. — If the tendency toward 
proximal fusion of polyps in Callozostron 
acanthodes is disregarded, the species can- 
not be aligned with any valid genus rec- 
ognized in the keys of Kiikenthal (1915, 
1919, 1924), and Bayer (1981). No genus 
having verticillate polyps with 8 marginal 
scales vertically aligned with the 8 oper- 
cular scales, as in Mirostenella, had been 
described. Although the sclerites of the 
polyps of Plumarella meet these qualifi- 
cations, the polyps of that genus are not 
arranged in whorls. In Mirostenella, all 8 
marginal scales have an acute apex or short 
spine; in Callozostron, as few as 4 or as 



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many as 8 marginal scales are prolonged 
as spines, but when all 8 are spinous (car- 
lottae, diplodiadema), the next one or two 
circles of body scales below the marginals 
also are spinous. 

Primnoella Gray 

Primnoella Gray, 1858:286; 1870:49.— 
Studer [& Wright], 1887:51 (part).— 
Wright & Studer, 1889:87 (part: B. "Car- 
inatae"). — Versluys, 1906:48, 52 (part: 
Carinatae). — Kukenthal 1907:210. — Au- 
rivillius, 1931:267. — Deichmann, 1936: 
162.— Bayer, 1956:F220 (part); 1981:936 
(part; in key only). — Broch, 1965:20 
(part). — Bayer & Stefani, 1989:455 (part; 
in key only). 

Primnoella "Compressae" Kukenthal, 
1908:13; 1912:311; 1915:148; 1919:384; 
1924:279. 

Type species. — Primnoa australasiae 
Gray, 1850, by monotypy. 

Diagnosis. — Flagelliform, unbranched or 
sparsely branched primnoids with polyps 
arranged in whorls, directed strongly up- 
ward, often appressed to axial cortex but 
never adnate; opercular scales decreasing in 
size toward axis, distinctly differentiated 
from, and more or less overreached by, 
marginal (i.e., "circumopercular") scales; 
body of polyps compressed, with two lon- 
gitudinal abaxial rows of wide imbricating 
scales, outer lateral and inner lateral rows 
reduced in size and numbers; adaxial sur- 
face either naked except for at most a few 
small scales below the marginals, or with 
small, delicate scales arranged in two more 
or less complete longitudinal rows. Surface 
of coenenchyme covered with rounded or 
polygonal scales; walls of longitudinal stem 
canals with irregular, tuberculate sclerites. 

Remarks. — The genus Primnoella Gray, 
1858, was based upon Primnoa austral- 



asiae Gray, 1850, syntypes of which are 
still preserved dry in The Natural History 
Museum, London (Reg. nos. BMNH 
1850.1.21.1, 6 and 10; 1983.3.2.13 and 14). 

Species groups within Primnoella sensu 
lato. — As heretofore defined, the genus 
Primnoella comprises two groups of spe- 
cies having polyps of distinctly different 
morphological types with respect to form 
and spiculation. In one group including the 
type species, designated "Carinatae" by 
Wright & Studer (1889:87) and later 
"Compressae" by Kiikenthal (1908:13), the 
polyps are directed strongly upward, more 
or less closely appressed to the coenenchy- 
me, with two longitudinal rows of large ab- 
axial scales, reduced outer and inner lateral 
scale rows, and their adaxial surface more 
or less naked and mostly devoid of sclerites 
below the marginals. In the other group, 
designated "Convexae," which includes P. 
magelhaenica, the polyps are curved in- 
ward less strongly if at all, there are four 
longitudinal rows of abaxial scales and well 
developed inner lateral and adaxial rows of 
scales so their adaxial surface is completely 
covered by sclerites. 

Although the polyps of some species of 
the Convexae group of Primnoella have 
fewer scales in the adaxial sclerite rows 
than in the abaxial rows, the adaxial side of 
the body nevertheless is completely covered 
by scales even if it is somewhat shorter than 
the abaxial side. There is no naked or nearly 
naked adaxial tract like that of all species 
of the Compressae group. Therefore, spe- 
cies of the Convexae group are separated 
from those of the Compressae group by a 
morphological discontinuity much more 
distinct and consistent than that separating 
many other related genus-group taxa in this 
and other families of Gorgonacea. This dis- 
continuity justifies the recognition of the 



Fig. 9. Callozostron carlottae Kukenthal, USNM 113%!. Part of one whorl. Top, Three polyps seen from 
below; Middle, Three polyps seen from above; Bottom, Detail of two polyps. Stereoscopic pairs (SEM 1627). 



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Convexae group of species as a genus dis- 
tinct from the Compressae group. 

Inasmuch as no genus-group name has 
heretofore been established for any species 
of the Convexae group, the name Convex- 
ella is now proposed for them as a distinct 
taxon, with Primnoella magelhaenica as 
type species. 

Distribution of Primnoella "Compres- 
sae." — The genus Primnoella as now re- 
stricted is most richly represented in Ant- 
arctic and sub-Antarctic waters, with rela- 
tively few species in temperate and tropical 
waters of the Atlantic, Pacific and Indian 
oceans. It is represented in the northern 
hemisphere by only one species, P. polita 
Deichmann, 1936. 

Species. — Kiikenthal (1924:280) as- 
signed 7 species to the "Compressae" 
group: P. scotiae Thomson & Ritchie, 
1906; compressa Kiikenthal, 1908; biseri- 
alis Wright & Studer, 1889; australasiae 
(Gray, 1850); delicatissima Kiikenthal, 
1909; and grandisquamis Wright & Studer, 
1889. To these may now be added P. polita 
Deichmann, 1936. As demonstrated below, 
Caligorgia (sic) gracilis Thomson & Mack- 
innon can also be assigned to Primnoella 
rather than to Callogorgia. 

Primnoella australasiae (Gray, 1850) 
Figs. 18, 19 

Primnoa australasiae Gray, T 850a: 146, Ra- 
diata pi. 2, figs. 8, 9; 1850b:510. 

Primnoella australasiae. — 1858:286; 1859: 
483; 1870:50.— Wright & Studer, 1889: 
88, pi. 18, figs. 1, la; pi. 21, fig. 15.— 
Versluys, 1906:52.— Thomson & Mack- 
innon, 1911:688, pi. 61, fig. 1.— Kiiken- 
thal, 1919:401, pi. 41, figs. 62, 63. 

Material examined. — D'Entrecaseaux 
Channel, between Tasmania and Bruce's Is- 
land. (Gray, teste Joseph Millingin, F. L. S. 



Syntypes, The Natural History Museum, 
London, register nos. 1850.1.21.1, 6 and 
10; 1983.3.2.13 and 14 (SEM 1936-1938). 
D'Entrecasteaux Channel (43°17'S, 
147°15'E), near Kinghorne Point, Bruny Is- 
land, Tasmania, 15 m, coll. A. J. Blackman, 
1979. One colony, USNM 58926 (SEM 
485, 487). Bluff Harbor, South Island, New 
Zealand (46°34'S, 168°20'E), coll. Dr. E. 
Kershner, January 1875. 30 colonies, dry, 
USNM 4505 (SEM 505). 

Discussion. — Robust, flagelliform colo- 
nies up to 1 m tall have 5-8 whorls of 13- 
15 strongly appressed polyps of the "Com- 
pressae" type in 1 cm of axial length. Only 
the two abaxial sclerite rows are fully vis- 
ible and contain 10-14 scales each; the out- 
er lateral rows each consist of about the 
same number of scales but they are smaller 
and because of their position only their edes 
are visible. The inner lateral rows each con- 
tain at least 5 or 6 scales, still smaller than 
those of the outer lateral rows. The walls of 
the longitudinal stem canals are filled with 
irregularly tuberculate sclerites of extreme- 
ly diverse shape. 

The dry specimens here reported from 
Bluff Harbor, New Zealand (Fig. 19, right) 
resemble Gray's type material (Fig. 18) 
from D'Entrecasteaux Channel, Tasmania. 
The colonies are attached by discoidal hold- 
fasts to the dead shells of pectinid bivalves, 
as many as 7 to a single shell, and reach a 
height of approximately 1 m. In some col- 
onies, the proximal 3 or 4 scales of the ab- 
axial rows are somewhat larger than those 
located more distally, and there is evidence 
that this may be related to the presence of 
eggs or embryos in the gastrovascular cav- 
ities. Although this condition was not no- 
ticed in the type specimens (Fig. 18), it was 
not specially looked for among the numer- 
ous dry specimens. 

The polyps of the topotypic colony from 



Fig. 10. Callozostron carlottae Kiikenthal, USNM 77387. Sclerites (SEM 1920). a. Opercular scales; b. 
Marginal scales; c. Body scales; d, Coenenchymal scales. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. IL Callozostron diplodiadema, new species, USNM 77386. Top, Complete whorl seen from above 
(SEM 1655); Bottom, Two whorls from the side (SEM 1628). Stereoscopic pairs. 



i 



VOLUME 109, NUMBER 1 



169 




Fig. 12. Callozostron diplodiadema, new species, USNM ll?i?i6. Top, Part of whorl, side view; Bottom, 
Detail of polyps. Stereoscopic pairs (SEM 1655). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 13. Callozostron diplodiadema, new species, USNM 77386. Detail of polyp. Top, Side view; Bottom, 
Oral view. Stereoscopic pairs (SEM 1656). 



Tasmania (Fig. 19, left) are somewhat more 
slender than those of either the type speci- 
mens examined or the colony from New 
Zealand and could be associated with the 



reproductive condition of the colony. The 
adaxial scales of this specimen preserved in 
alcohol seem not so closely fitted as in the 
two lots of dried specimens. 



VOLUME 109, NUMBER 1 



171 



The specimens sent to Gray were "cov- 
ered with a calcareous coat of a cream-yel- 
low colour" (Gray, 1850:147). The illustra- 
tion given by Thomson & Mackinnon 
(1911; pi. 61, fig. 1) gives a fair idea of the 
appearance of the whorls. 

Primnoella laevis 

(Thomson & Mackinnon, 1911) 

Fig. 20 

Caligorgia laevis Thomson & Mackinnon, 
1911:689, pi. 65, fig. 1; pi. 68, fig. 7; pi. 
80.— Kiikenthal, 1919:377; 1924:275, 
fig. 155. 

Callogorgia laevis. — Bayer, 1982:122 (in 
key only; removed from Callogorgia and 
referred to Primnoella in footnote, but 
new combination was not actually printed 
there). 

Material examined. — Syntypes (Austra- 
lian Museum, Sydney). 

Description. — See Thomson & Mackin- 
non, 1911:689. 

Distribution. — Coast of New South 
Wales. 

Remarks. — While on a brief visit to the 
Australian Museum, I was able to examine 
Thomson & Mackinnon's original material 
through the kindness of Dr. Pat Hutchings. 
As can be seen from the photograph of a 
specimen reproduced by Thomson & 
Mackinnon (1911:pl. 80), the branching of 
the colony is spare and open rather than 
"luxuriant" as described by Thomson & 
Mackinnon. Even the rather crude drawing 
of a single polyp given by Thomson & 
Mackinnon (1911:pl. 55, fig. 1) is adequate 
to demonstrate that the species does not be- 
long to ''Caligorgia'" (= Callogorgia). Al- 
though the polyps (Fig. 38) are arranged in 
whorls as usual in both Callogorgia and 
Primnoella and are directed obliquely up- 
ward and curved toward the coenenchyme, 
they do not face inward toward the axis as 
is usual in Callogorgia, and the body scle- 
rites are not strongly sculptured externally 
as commonly is the case in Callogorgia 
(e.g., see Bayer 1982:figs. 2, 4). 



Although Primnoella divaricata (Studer) 
and P. divergens Hickson are described as 
branched they are very sparsely so, whereas 
P. laevis is repeatedly branched in a di- 
chotomous manner. 

Convexella, new genus 

Primnoella. — Studer [& Wright], 1887: 
51. — Madsen, 1944:39.— Bayer, 1956: 
F220 (part).— Broch, 1965:20 (part).— 
Bayer, 1981:936 (part; in key only). — 
Bayer & Stefani, 1989:455 (part; in key 
only). 

Primnoella Convexae. — Wright & Studer, 
1889:83.— Versluys 1906:51.— Kiiken- 
thal, 1908:13; 1912:311; 1915:148; 1919: 
384; 1924:279. 

Type species.— Primnoella magelhaeni- 
ca Studer, 1879, here designated. 

Diagnosis. — Flagelliform, unbranched or 
sparsely branched primnoids with polyps 
arranged in whorls, standing almost verti- 
cally or directed more or less obliquely up- 
ward, not appressed to axial cortex; mar- 
ginal and sometimes submarginal scales 
folding over opercular scales, which are 
more or less hidden and not clearly differ- 
entiated from marginals; body of polyps cy- 
lindrical and weakly curved, not com- 
pressed, abaxial surface covered by four 
longitudinal rows of rounded or oval im- 
bricating scales, adaxial and lateral surfaces 
covered by inner-lateral and adaxial scales 
of similar size, in rows that may be some- 
what shorter than the abaxial and outer-lat- 
eral rows. Surface of coenenchyme covered 
with rounded, imbricating scales; walls of 
longitudinal stem canals with irregular, tu- 
berculate sclerites. 

Remarks. — The type species is now il- 
lustrated extensively to demonstrate its 
variability and to show its similarities to 
Callozostron, without prematurely speculat- 
ing upon phylogenetic relationships. 

Species. — Kukenthal (1924:280) referred 
seven species to the "Convexae" group of 
Primnoella: flagellum Studer, 1879; van- 
hoejfeni Kukenthal, 1909; magelhaenica 



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VOLUME 109, NUMBER 1 



173 




Fig. 15. Callozostron acanthodes, new species, USNM 94575. Top, Part of branch with two whorls of 
polyps; Bottom, Detail of whorl. Stereoscopic pairs (SEM 1594). 



Fig. 14. Callozostron diplodiadema, new species, USNM 77386. Sclerites (SEM 1657). a. Opercular scales; 
b, Marginal and submarginal scales; c. Body scales with acute apex; d. Body scales; e, Coenenchymal scales. 



174 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




159409 10KV; X80.0 .38mm 



159411 10KV K80.0 .38mm 



Fig. 16. Callozostron acanthodes, new species, USNM 94575. Top, Part of branch viewed from above, 
showing two pairs of basally united polyps (SEM 1595); Bottom, Distal end of polyp showing opercular scales 
in closed position (SEM 1594). Stereoscopic pairs. 



VOLUME 109, NUMBER 1 



175 




Fig. 17. Callozostron acanthodes, new species, USNM 94573. 
Marginal scales; c. Opercular scales; d, Coenenchymal scales. 



Sclerites (SEM 1618). A, Body scales; b. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 18. Primnoella australasiae Gray, BM(NH) 1850.1.21.1/6/10. Isolated polyp in abaxial, lateral, and 
oral view. Stereoscopic pairs (SEM 1936). 



VOLUME 109, NUMBER 1 



177 



..- i. 









' ' 'St' 



.V 



^i\ 



^ 



^An'^, 





U\k:^hA 



'^xVju^^^xLiL.A. '<■'• het^jtSd' 



Fig. 19. Primnoella australasiae Gray. Left, from D'Entrecasteaux Channel, Tasmania, USNM 58926; Right, 
from Bluff Harbor, New Zealand, USNM 4505. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 20. Primnoella laevis (Thomson and Mackinnon), from east of Broken Bay, N.S.W., Australia. Two 
whorls from syntype, scale bar = 0.5 mm. Stereoscopic pairs (SEM 772). 



Studer, 1879; murrayi Wright & Studer, 
1889; antarctica Kiikenthal, 1907; distans 
Studer, 1879; divaricata (Studer, 1879); and 
diver gens Hickson, 1907. 

Of these, Primnoella flagellum Studer, 
1879, is here interpreted as a variant of P. 
magelhaenica Studer, 1879. 

Kiikenthal (1912:317) described Prim- 
noella vanhoejfeni as similar to P. magel- 



lanica (sic), but his illustrations are not 
convincing. 

Primnoella murrayi Wright & Studer 
(1889:84, pi. 18, figs. 3, 3a) is similar to P. 
magelhaenica but has a short apical spine 
or acute angle on the distal margin of the 
abaxial body scales, a character so variable 
in P. magelhaenica that the species can be 
considered synonymous. 



VOLUME 109, NUMBER 1 



179 



Kiikenthal (1919:387, 391) placed his 
Primnoella antarctica in the "Convexae" 
group of species because the polyps are not 
compressed, even though the adaxial side is 
mostly naked except for a few small scales 
below the mouth and near the base; it bears 
some resemblance to Ophidiogorgia but 
none at all to Convexella magelhaenica. 

The polyps of Primnoella distans Studer, 
1879, have only two visible rows of large 
abaxial scales (Wright & Studer 1889, pi. 
18, fig. la), and very small, reduced adaxial 
scales, so must be referred to the "Com- 
pressae" group of species. 

Based upon Studer's type, Versluys 
(1906:54) described Primnoella divaricata 
(Studer, 1879), as very similar to P. austra- 
lasiae but with the adaxial side of the pol- 
yps completely covered by scales smaller 
than those of the abaxial rows; it conforms 
better with the "Compressae" group and 
therefore is here treated as a species of 
Primnoella. 

About all that can be said about Prim- 
noella divergens, given the inadequate de- 
scription and figure originally provided 
(Hickson, 1907:10, pi. 1, figs. 8-10), is that 
it is not one of the "Compressae" group 
but may be a species of "Convexae." 

As originally described and illustrated, 
both Primnoella jungerseni Madsen, 1 944, 
and P. krampi Madsen, 1956, fall within the 
"Convexae" group of species, hence are 
here reassigned to Convexella. Although 
Madsen (1944:42) compared P. jungerseni 
with P. distans and P. polita Deichmann, 
1936, in both of those species only two 
rows of abaxial scales are visible, making 
both referable to the "Compressae" group, 
hence Primnoella s.s. 

Primnoella gracilis Molander, 1929, 
originally was compared with P. flagellum 
and, as described and illustrated (Molander 
1929:63, figs. 17, 18; pi. 1, fig. 2), falls 
within the "Convexae" group on the basis 
of its well-developed adaxial scales. How- 
ever, the marginal scales show no tendency 
to override the operculars, as is the case in 
both Primnoella and Convexella, so the 



species probably is excluded from both 
genera. 

Gravier (1914:77) aligned his Primnoella 
kuekenthali with the "Convexae" group 
(1914:83), but his illustrations and lengthy 
description throw some doubt upon that as- 
signment. The body scales are thick, scup- 
tured externally by large ridges that may 
branch, and have strongly dentate margins, 
whereas the sclerites of Primnoella "Con- 
vexae" are thin and externally smooth. Gra- 
vier's statement that on the concave adaxial 
surface of the polyps "il y a une bande me- 
diane qui reste a nu" suggests that kuek- 
enthali may be referable to Ophidiogorgia 
Bayer, 1980. This conclusion is confirmed 
by specimens from Marguerite Bay 
(68°30'S, 68°30'W, USNM 56631), South 
Shetland Islands (62°17'30"S, 58°34'36"W, 
USNM 78398; 63°26'S, 62°10'W, USNM 
77456; 62°16'42"S, 58°34'00"W, USNM 
77361), Livingston Island (63°24'S, 62°14'W, 
USNM 82863), Anvers Island (64°46'28"S, 
63°26'30"W, USNM 81544), Elephant Island 
(62°39'S, 56°10'W, USNM 82852), Adelaide 
Island (67°48'12"S, 68°54'06"W, USNM 
77460), all from along the Antarctic Penin- 
sula in the general vicinity of the type locality 
(64°50'S, 63°30'W), as well as from the 
South Orkney Islands (60°26'30"S, 
45°53'18"W, USNM 77145), that agree in all 
essentials with Gravier's description of Prim- 
noella kuekenthali. As in his material, the in- 
terval between whorls and the size of polyps 
within whorls is subject to considerable vari- 
ation; in two specimens (USNM 56631, 
77456), new whorls are intercalated between 
fully developed whorls just as illustrated by 
Gravier (1914:82, fig. 108). Gravier's draw- 
ing shows the body scales aligned longitudi- 
nally, unlike the case in Ophidiogorgia par- 
adoxa Bayer but, in the specimens now avail- 
able, this regular arrangement is disrupted in 
almost all fully developed polyps. It must be 
remembered that the then unique type speci- 
men of O. paradoxa was deformed by the 
presence of some epizoan, possibly echino- 
derm, which may have disrupted the devel- 
opment of sclerites in the polyps of adjacent 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 21. Convexella magelhaenica (Studer), USNM 85300, from north of Cabo San Diego, Tierra del Fuego, 
124—119 m, showing variation present among specimens from a single haul. See Figs. 27, 28. 



whorls. Although Gravier described the dis- 
talmost scales of the polyps as an operculum, 
his drawing (1914:81, fig. 105) of one oper- 
cular scale is no more clearly differentiated 
from the body scales than are those of O. 
paradoxa. Accordingly, Primnoella kueken- 
thali Gravier, 1913, is here transferred to 
Ophidiogorgia as O. kuekenthali, new com- 
bination, pending detailed studies to clarify 
its relationship with O. paradoxa. 

Distribution. — Except for Convexella 
jungerseni (Madsen), all species of Convex- 
ella are confined to the southern hemi- 
sphere. 

Specimens from Balleny Islands (66°53'S, 
163°19'E, USNM 77330) and the vicinity of 
Wilkes Station (66°15'24"S, 110°28'40"W, 
USNM 88775) clearly referable to Ophidi- 
ogorgia indicate that the genus and probably 



the species are of circum-Antarctic distribu- 
tion. 

Convexella magelhaenica 

(Studer, 1879), new combination 

Figs. 21-38 

Primnoella magelhaenica Studer, 1879:644, 
pi. 2, fig. lOa-c— Kukenthal, 1919:389, 
pi. 40, figs. 50, 51; 1924:281 (refer- 
ences). — Broch, 1965:21, pi. 1, fig. 1. 

Primnoella magellanica Wright & Studer, 
1889:83, pi. 17, fig. 2; pi. 21, fig. 10. (in- 
correct subsequent spelling). — Thomson 
& Ritchie, 1906:855, pi. 1, fig. 3 (incor- 
rect subsequent spelling). — Versluys, 
1906:51 (unjustified emendation). 

Primnoella flagellum Studer, 1879:645, pi. 
2, fig. lla-c— Wright & Studer, 1889: 



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181 




Fig. 22. Convexella magelhaenica (Studer). Left, Colony from off Isla Desolacion, Magellanes, Chile, 1500- 
1666 m, showing polyps directed distad on distal part, basad on basal part, USNM 82856 (see Fig. 35 for detail 
of polyps); Right, Two colonies from off Cabo San Sebastian, Tierra del Fuego, 86 m, USNM 85306, showing 
polyps standing vertically as in Callozostron (compare figs. 1, right, and 7, top; see Figs. 23, 24 for detail of 
whorls and polyps). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 23. Convexella magelhaenica (Studer), USNM 85306, from off Cabo San Sebastian, Tierra del Fuego, 
86 m. Top, Three whorls; Bottom, Detail of polyps showing development of apical spines on distal body scales. 
Stereoscopic pairs (SEM 1644). 



85, pi. 18, figs. 2, 2a; pi. 21, fig. 12.— 
Versluys, 1906:51. 
IPrimnoella flagellum. — Thomson & 
Mackinnon, 1911:688. 



Material examined. — 35 lots. See appen- 
dix. 

Discussion. — Studer's (1879) original 
description and illustrations of P. magel- 



VOLUME 109, NUMBER 1 



183 




Fig. 24. Convexella magelhaenica (Studer), USNM 85306, from off Cabo San Sebastian, Tierra del Fuego, 
86 m. Top, Whorl of polyps seen from above; Bottom, Oblique view of polyp showing circumoperculum. 
Stereoscopic pairs (SEM 1645). 



haenica, collected at a depth of 42 fathoms 
(= 77 m) in the Straits of Magellan by the 
Gazelle, are not adequate to distinguish the 
species from others described subsequently. 



Wright & Studer (1889), Thomson & Rit- 
chie (1906), Kukenthal (1919), and Broch 
(1965) have described and illustrated spec- 
imens purported to be magelhaenica, but in 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 25. Convexella magelhaenica (Studer), USNM 89344, from between Tierra del Fuego and Falkland 
Islands, 128 m. Top, Part of slender colony with whorl of new polyps between fully developed whorls (SEM 
1934); Bottom, Part of stouter colony showing new polyp between fully developed individuals within a whorl, 
as well as new whorl almost hidden by fully developed whorl (SEM 1931). Stereoscopic pairs. 



VOLUME 109, NUMBER 1 



185 




Fig. 26. Convexella magelhaenica (Studer), USNM 89344, from between Tierra del Fuego and Falkland 
Islands, 128 m. Part of stout colony with polyps exceptionally crowded. Stereoscopic pairs (SEM 1930). 



no case was identification confirmed by 
comparison with the type specimen. It is 
reasonably certain that the specimens in the 
present collection are conspecific with 
Thomson «& Ritchie's, Kiikenthal's, and 
Broch's material. Wright & Studer's de- 
scriptions and illustrations of both P. ma- 
gellanica (incorrect subsequent spelling of 
magelhaenica) and P. flagellum are consis- 
tent with the numerous specimens now on 
hand from the same general geographical 
area. It should be remembered, however, 
that if future study should prove Wright & 



Studer's specimen of ''magellanica" to rep- 
resent a species different from P. magel- 
haenica, it may take the available name ma- 
gellanica dating from Versluys, 1906. 

The present material, consisting of spec- 
imens of a wide range in size, from young 
colonies 2.5 cm tall with 7 whorls of polyps 
in 2's and 3's, to mature specimens more 
than 40 cm tall, with 7 whorls of 15 polyps 
each in 1 cm, comprises colonies agreeing 
with the characters heretofore described for 
P. magelhaenica, P. flagellum, and P. ''ma- 
gellanica." Colonies of intermediate size 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 27. Convexella magelhaenica (Studer). Top, USNM 85300, from north of Cabo San Diego, Tierra del 
Fuego, 124-1 19 m; part of slender colony with whorl of new polyps between two fully developed whorls (SEM 
1926). Bottom, USNM 89344, from between Tierra del Fuego and Falkland Islands, 128 m; part of stouter 
colony with crowded whorls of different ages (SEM 1932). Stereoscopic pairs. 



VOLUME 109, NUMBER 1 



187 




Fig. 28. Convexella magelhaenica (Studer), USNM 85300, from north of Cabo San Diego, Tierra del Fuego, 
124—119 m. Top, Part of stout colony with whorl of upturned new polyps between two fully developed whorls 
of outwardly directed polyps (SEM 1929); Bottom, Part of stouter colony having polyps with exceptionally 
strong circumopercular spines (SEM 1928). Stereoscopic pairs. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 29. Convexella magelhaenica (Studer), USNM 58441, from Strait of Le Maire, 229-265 m. Top, Part 
of branch showing young whorl with polyps turned strongly upward, situated between fully developed whorls 
with some polyps projecting almost vertically from axis; Bottom; Detail of two polyps. Stereoscopic pairs (SEM 
1629). 



may have from 3 to 7 whorls in 1 cm of 
axial length, varying from 3 to 8 or more 
polyps per whorl, the distalmost sometimes 
only 2. 

Broch (1965:21) did not provide a formal 



decription of his specimens of P. magel- 
haenica from Burdwood Bank but com- 
mented that his "rather copious collection 
exhibits a comparatively broad variation" 
from "slender specimens" to "more coarse 



VOLUME 109, NUMBER 1 



189 




Fig. 30. Convexella magelhaenica (Studer), USNM 58442, from east of Cape Horn, 384-394 m. Part of 
stem with polyps bent upward but not appressed to stem; middle portion in stereoscopic view, scale bar = 1 
mm (SEM 389). Isolated sclerites (SEM 1922): top row, operculars; second row, outer (left group) and inner 
(right group) coenenchymals; bottom row, circumopercular and body scales. 




PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



V 




0.5KV 



S.dSmm 






0.5KV 



2.09mn) 




1647 0.5KV 



. ?5inm 



1647 0.5KV 



. 75mm 



Fig. 31. Convexella magelhaenica (Studer), USNM 85288, from Sars Seamount in Drake Passage, 512-622 
m. Top, Part of branch with whorls of polyps; Bottom, Polyp in side view, showing negligible development of 
marginal points on distal body scales. Stereoscopic pairs (SEM 1647). 



colonies." His specimens were 12-18 cm 
long, a size not unusual in the material re- 
ported here. However, the longest specimen 
from Eltanin station 217 from off Cabo San 



Diego, Tierra del Fuego (USNM 78403), is 
a slender colony 48 cm long. The type spec- 
imen of Primnoella magelhaenica was 22 
cm long, and that of P. flagellum 53 cm, so 



VOLUME 109, NUMBER 1 



191 




Fig. 32. Convexella magelhaenica (Studer), USNM 85288, from Sars Seamount in Drake Passage, 512-622 
m. Top, Polyp in abaxial view; Bottom, Oral view of polyp with circumopercular scales closed over operculars. 
Stereoscopic pairs (SEM 1647). 



it is probable that the former is a colony 
similar to the "more coarse colonies" men- 
tioned Broch and abundantly represented in 
the present material, whereas P. flagellum 
is like Broch's "slender specimens." Both 



extremes are present in the lots from Eltan- 
in stations 217 (USNM 78403), 369 
(USNM 77371), 974 (USNM 85300), and 
976 (USNM 89344), all from east of Tierra 
del Fuego. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 33. Convexella magelhaenica (Studer), USNM 85298, from Strait of Le Maire, 641-596 m. Top, Detail 
of whorl showing polyps in abaxial and lateral views (SEM 1648); Bottom, Young polyps developing between 
adult polyps within two whorls from which some of the adult polyps have been removed. The young polyps 
have 7 scales in the abaxial rows compared to 13 in the adults, showing that the number of scales can be age 
dependent (SEM 1649). Stereoscopic pairs. 



VOLUME 109, NUMBER 1 



193 



Broch (1965:20-22) commented on the 
production of new polyps and new whorls 
in Primnoella and reported some differ- 
ences in this respect among P. magelhaen- 
ica, P. scotiae, and P. vanhoejfeni. He re- 
ported that new whorls arise in the middle 
and upper parts of the polyparium in P. ma- 
gelhaenica. In the present material, devel- 
opment of new whorls is very variable. In 
some colonies, few new whorls are inter- 
calated between fully developed whorls but, 
in others, new whorls may arise throughout 
the length of the colony, indicating different 
states of colonial growth at the time of col- 
lection. In a specimen from Eltanin station 
976, two new whorls of different sizes may 
occur in the space between two fully de- 
veloped whorls. 

Studer (1879:645) reported that the body 
scales "stehen in 13-14 Querreihen um die 
Kelche," and Wright & Studer (1889:83) 
wrote that "ten to thirteen calyx scales may 
be counted" in the length of the calyx, but 
in neither case do the accompanying illus- 
trations suggest so many. Studer (1879:645) 
said that in P. flagellum the body scales 
"stehen um die Kelche in acht Reihen," a 
number that occurs in the present material. 

Colonies are sometimes attached to a 
stone or shell by a small, discoidal, calcified 
holdfast, but the supporting substrate is not 
often collected with the specimens. Most 
are broken off just above the holdfast, part 
of which may be present, suggesting that 
the usual substrate is rocky. However, the 
numerous slender colonies obtained at El- 
tanin station 976 show no sign of attach- 
ment. Although all specimens are more or 
less broken and the axis of most is partially 
decorticated, in no case can a growing tip 
at both ends be confirmed. Pogonophoran 
tubes among the gorgonian stems suggest 
that the bottom was muddy at this location, 
making it plausible to conclude that the 
very flexible colonies lie prone on the sub- 
strate in life, as is the case with some spe- 
cies of Leptogorgia (Bayer 1961:218-222, 
Grasshoff 1988:116). 

The straight or slightly curved polyps 



vary in their stance from almost perpendic- 
ular to the axis, to directed strongly upward. 
In no case are they strongly bent inward 
with the operculum facing the axis. A spec- 
imen without holdfast (USNM 82856, Fig. 
21, left) has the polyps along the distal half 
of the colony directed toward the apex, 
those along the proximal half toward the 
base. In any given colony, the number of 
body scales in the abaxial rows may vary 
from 7 or 8 in polyps near the middle of 
the colony, to as few as 5 in the polyps of 
the distalmost and basalmost few whorls; in 
colonies with the tallest polyps, the abaxial 
rows commonly consist of 10-1 1 scales be- 
low the circumoperculars, but as many as 
14 is unusual. Young polyps have fewer ab- 
axial scales than do fully developed indi- 
viduals (Figs. 25, 27, 33). 

Close observation of isolated polyps dur- 
ing maceration with Sodium hypochlorite 
shows that the scales of the distalmost cir- 
cle of 8 body sclerites forming the circu- 
moperculum are not distinctly differentiated 
functionally from the following 2 or 3 cir- 
cles, all of which participate in protecting 
the mouth and withdrawn tentacles during 
contraction. The distalmost narrow, tongue- 
shaped sclerites (Fig. 30) that comprise the 
theoretical "operculum" can fold nearly 
flat over the mouth in contraction. These 
are not consistently 8 in number but vary 
from 6 to 9 or even 10 among polyps of 
the same whorl. In some cases these differ 
in size among themselves only slightly and 
are distinctly smaller than the circumoper- 
cular scales; in others the adaxials are very 
small and the abaxials are as large as, and 
indistinguishable from, the circumopercu- 
lars. These and intermediate conditions ex- 
ist not only in a single colony, but even 
among the polyps of one whorl. 

The arrangement of opercular and cir- 
cumopercular sclerites resembles that de- 
scribed for Thouarella hilgendorfi by Ki- 
noshita (1908:5, fig. 1). In both Thouarella 
and Primnoella, sclerites are so large that 
eight scales cannot fit around the circum- 
ference of the polyp in a single circle. In- 



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VOLUME 109, NUMBER 1 



195 



Stead, each circle of eight scales surround- 
ing the body is composed of two "trans- 
verse rows" of four scales that alternate 
with one another (Kinoshita 1908), so the 
"operculum" consists of transverse rows I 
and II, and the circumoperculum of rows III 
and IV. The scales seem to be arranged in 
spirals around the polyp as well as longi- 
tudinally, as is shown in Fig. 23. 

The abundant material here reported 
strongly suggests that Studer's Primnoella 
magelhaenica and P. flagellum represent a 
single species, and that Wright & Studer's 
Kiikenthal's, and Broch's records of "ma- 
gellanicd" and magelhaenica are the same 
thing. The present material comes from the 
remarkable bathymetric range from 75 to 
more than 1500 m. The general aspect of 
some colonies from the two bathymetric ex- 
tremes is considerably different and would, 
in the absence of intermediates, easily be 
considered specifically distinct. However, 
morphological variation among specimens 
from the extremes of bathymetric range dif- 
fers little from that present among speci- 
mens from a single shallow station, making 
discrimination of two or more species im- 
possible on any objective basis so far de- 
tected. 

To convey some idea of the similarities 
and differences among colonies of P. ma- 
gelhaenica over its wide bathymetric range, 
scanning stereomicrographs have been 
made of a selection of specimens from the 
shallowest to the deepest available records, 
some of which are here presented to docu- 
ment the morphological diversity of this 
abundant gorgonian. 

The shallowest record in the present col- 
lection is 73 m, obtained off the north en- 
trance of the Strait of Lemaire, Tierra del 
Fuego (USNM 85308). Specimens from 86 



m off Cabo San Sebastian (USNM 85306) 
are typical of shallow-water colonies (Figs. 
23, 24). The polyps are directed obliquely 
upward but the whorls do not overlap in the 
contracted condition; scales of the three dis- 
talmost rings of body sclerites have a dis- 
tinct, smooth apical spine, and the theoret- 
ical operculum is concealed during contrac- 
tion by the uppermost rings of circumoper- 
cular scales. 

Colonies trawled from 128 m between 
Tierra de Fuego and the Falkland Islands 
(USNM 89344) reveal a range of growth- 
form from slender, with well separated 
whorls of few polyps (Fig. 25, top) to un- 
usually stout, with crowded, overlapping 
whorls of eight or more polyps (Fig. 26). 
Small, evidently young polyps may occur 
between fully developed individuals within 
a whorl (Fig. 25, bottom). 

A haul made north of Cabo San Diego, 
Tierra del Fuego in 124-1 19 m yielded sev- 
eral colonies of diverse aspect (USNM 
85300), from slender (Fig. 27, top) to rather 
stout (Fig. 28). New whorls are intercalated 
between fully developed ones (Figs. 27, 
top; 28, top), and the younger individuals 
may be more strongly inclined toward the 
axis than are the older (Fig. 28, top), which 
may stand almost vertically. In one colony 
the circumopercular scales are so strongly 
developed that it approaches Callozostron 
in appearance (Fig. 28, bottom). 

Colonies from the Strait of Le Maire in 
229-265 m (USNM 58441) demonstrate 
the insertion of new whorls of polyps be- 
tween fully developed whorls of almost 
perpendicularly placed individuals (Fig. 
29). 

A colony from east of Cape Horn in 384- 
494 m (USNM 58442) has the tallest pol- 
yps observed in the present material (Fig. 



Fig. 34. Convexella magelhaenica (Studer), USNM 82865, from south of Isla de los Estados, Tierra del 
Fuego, 771-903 m. Top, Part of branch showing two whorls of polyps with points scarcely developed on distal 
body scales; Middle, Oblique view of polyps showing scant development of marginal points on body scales; 
Bottom, Oral view of polyp with circumopercular scales in closed position. Stereoscopic pairs (SEM 1651). 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 35. Convexella magelhaenica (Studer), USNM 82856, from off Isla Desolacion, Magellanes, Chile, 
1500—1666 m. Top, Part of stem with two whorls; Bottom, Polyp in side view, showing alternating tall and wide 
circumopercular scales and weak development of apical points. Stereoscopic pairs (SEM 1643). 



30). They have 12-13 scales in the abaxial 
rows, and only the scales of the distalmost 
transverse row have a strong apical point. 
The polyps of a colony from Sars Sea- 



mount in Drake Passage trawled in 512- 
622 m (USNM 85288) are strongly inchned 
upward and have as many as 13 or 14 scales 
in the abaxial rows; the the circumopercular 



VOLUME 109, NUMBER 1 



197 




Fig. 36. Convexella magelhaenica (Studer), USNM 85302, from east edge of Burdwood Bank, 1647-2044 
m. Top, Part of stem with three whorls; Bottom, Detail of polyps. Stereoscopic pairs (SEM 1652). 



scales completely obscure the operculars, 
and have only a low, rather blunt apex 
(Figs. 31, 32). 

A colony from 641-586 m in the Strait 
of Le Maire (USNM 85298) has polyps 



with 10-11 scales in the abaxial rows and 
circumopercular scales with weakly devel- 
oped apical angle (Fig. 33). Young polyps 
developing between fully developed indi- 
viduals have fewer abaxial scales, showing 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 37. Convexella magelhaenica Stxider, USNM 59102, from east of Cape Horn, 384-494 m. Two whorls; 
camera lucida drawing by Constance Stolen McSweeny from alcoholic specimen, for contrast with SEM images. 
The tubercles on the inner surface of the scales are clearly visible through the glassy clear sclerites, which are 
opaque and externally smooth as depicted by SEM. 



that the number of scales in the longitudinal 
rows increases with age. 

In colonies from depths of 771-903 m 
south of Isla de los Estados, Tierra del Fue- 
go (USNM 82865), the uppermost four 
scales of the circumoperculum are tall and 
triangular, alternating with the lower four 
scales which are broader and have a low 
apical point (Fig. 34). 



Colonies from west of Isla Desolacion, 
Chile, in 1500-1666 m (USNM 82856) are 
slender, with well spaced whorls of 5 or 6 
upwardly directed polyps (Fig. 35). Four 
tall, triangular circumopercular scales alter- 
nate with four broader scales with a low 
apical angle. 

The tendency to be slender and more del- 
icate continues in colonies from 1647-2044 



VOLUME 109, NUMBER 1 



199 




Fig. 38. Convexella magelhaenica (Studer), USNM 59102, from east of Cape Horn, 384-494 m. SEM image 
for comparison with camera lucida drawing in Fig. 36. (SEM 1646.) 



m on the eastern edge of Burdwood Bank 
(USNM 85302), which have well spaced 
whorls of 3 or 4 upwardly directed polyps 
(Fig. 35). The four lower circumopercular 
scales are shorter than the four upper scales, 



but are more like them than is the case in 
USNM 82856. 

Remarks. — Because heavy reliance upon 
scanning electron microscopy for highly ac- 
curate representation of external morphol- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ogy causes us to lose sight of the fact that 
primnoid sclerites are more or less translu- 
cent, a camera lucida drawing (Fig. 37) of 
one specimen is given for comparison with 
SEM representation (Fig. 38). The sclerites 
of P. magelhaenica are translucent, almost 
glassy clear when wet, so the tubercular 
sculpture of their inner surfaces is clearly 
visible with the light microscope but invis- 
ible with SEM. 

In view of the morphological similarities 
of Primnoella magelhaenica with Callozos- 
tron carlottae Kiikenthal and C diplodi- 
adema new species, it might be as reason- 
able either to regard P. magelhaenica as a 
weakly spinose Callozostron, or C. carlot- 
tae and C. diplodiadema as very spiny spe- 
cies of Primnoella, apart from the fact that 
irregularly tuberculate sclerites are abun- 
dantly present in the walls of the longitu- 
dinal stem canals in P. magelhaenica as 
they are in P. australasiae, but not in spe- 
cies of Callozostron. 

Distribution. — West and east coasts of 
the southern extremity of South America, 
Straits of Magellan and Le Maire, Drake 
Passage, Tierra del Fuego, eastward to 
Falkland and South Orkney Islands, 73- 
1647 m (this collection). Burdwood Bank, 
61 fathoms (Broch 1965). This species has 
not been found in any of its variants any- 
where else around the Antarctic continent. 

Acknowledgments 

Dr. P. F S. Cornelius of The Natural His- 
tory Museum (London) made it possible for 
me to examine Gray's type specimens of 
Primnoella australasiae, and Dr. D. A. Ritz 
provided specimens recently collected in 
the type locality by A. J. Blackman. Dr. Pat 
Hutchings of the Australian Museum (Syd- 
ney) provided access to Thomson & Mack- 
innon's type material of Caligorgia laevis. 
Dr. Manfred Grasshoff of the Naturmuseum 
Senckenberg (Frankfurt) generously shared 
his broad knowledge of gorgonacean tax- 
onomy during the preparation of this manu- 
script and provided many helpful criticisms. 



The numerous scanning electron micro- 
graphs that vastly enhance the value of this 
paper were made by Mr. Walter R. Brown, 
head of the SEM Laboratory, National Mu- 
seum of Natural History. Plates illustrating 
the sclerites were assembled and mounted 
by Ms. Molly Ryan, staff illustrator. De- 
partment of Invertebrate Zoology, National 
Museum of Natural History. I here express 
my sincere thanks to all those who contrib- 
uted their time and skills toward the prep- 
aration of this report. 

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Appendix 

Convexella magelhaenica (Studer) 

Material examined (listed in sequence of USNM 
catalogue numbers). 

58441: Strait of Le Maire: 54°56'S, 65°03'W, 229- 
265 m, Eltanin sta. 969, 10 Feb 1964. Two colonies 
with holdfast, one attached to bivalve shell. (SEM 
1629.) 

58442: East of Cape Horn: 56°06'S, 66°19'W, 384- 
494 m, Eltanin sta. 740, 18 Sep 1963. One colony. 
(SEM 389.) 

59102: East of Cape Horn: 56°06'S, 66°19'W, 384- 
494 m, sta. 740, 18 Sep 1963. Four small colonies 
originally attached to pebbles and shell, now detached. 
(SEM 1646.) 

59288: East of Cape Horn: 56°06'S, 066°19'W, 384- 
494 m, Eltanin sta. 740, 18 Sep 1963. One colony. 

77371: Tierra del Fuego, east northeast of Cabo San 
Diego: 54°04'S, 63°35'W, 247-293 m, Eltanin sta. 
369, 12 Dec 1962. 25 colonies, more or less damaged, 
and 3 denuded axes. 

77394: East edge of Burdwood Bank: 54°43'S, 
55°30'W, 1647-2044 m, Eltanin sta. 1592, 14 Mar 
1966. 16 colonies, 8 attached to stones, 5 with part of 
holdfast, 3 lacking holdfast. (SEM 1652, 1653, 1924.) 

78399: Tierra del Fuego, off north entrance to Strait 
of Le Maire: 54°29'S, 64°00'W, 104 m, RA^ Hero sta. 
863, 22 Oct 1971. Five colonies, one badly damaged. 

78403: Tierra del Fuego, east northeast of Cabo San 
Diego: 54°22'S, 64°42'W, 106-110 m, Eltanin sta. 
217, 23 Sep 1962. Five colonies lacking holdfast. 

78656: Tierra del Fuego, north of Isla de los Esta- 
dos: 54°50'S, 63°59'W, 0205-0208 m, RA^ Hero sta. 
880, 28 Oct 1971. One colony lacking holdfast. 

78657: Tierra del Fuego, north of Isla de los Esta- 
dos: 54°34'S, 64°40'W, 84-85 m, RA^ Hero sta. 903, 
6 Nov 1971. One colony lacking holdfast. 

78661: Chile, west of Isla Desolacion, Magellanes: 
52°52'S, 75°18'W, 119-329 m, Eltanin sta. 288 Cruise 
21*, 6 Dec 1965. Four colonies lacking holdfast, and 
three fragments. 

82847: Between Tierra del Fuego and Falkland Is- 
lands: 52°45'S, 66°34"W, 92 m, Eltanin sta. 337, 2 Dec 
1962. Two damaged colonies. 

82856: Chile, west of Isla Desolacion, Magellanes: 



53°13'S, 75°41'W, 1500-1666 m, Eltanin sta. 283 
Cruise 21*, 5 Dec 1965. Three colonies, two some- 
what damaged. (SEM 1643, 1923.) 

82865: Tierra del Fuego, south of Isla de los Esta- 
dos: 54°55'S, 64°00'W, 0771-0903 m, RA^ Hero sta. 
875, 27 Oct 1971. Three colonies, smallest with hold- 
fast, and three denuded axes attached to shells. (SEM 
1651.) 

82866: Tierra del Fuego, off Isla de los Estados: 
54°46.9'S, 64°04'W, depth not recorded, RfV Hero sta. 
881, 29 Oct 1971. Two damaged colonies. 

82867: Tierra del Fuego, north of Isla de los Esta- 
dos: 54°34'S, 64°30'W, 73-75 m, R/V Hero sta. 907, 
7 Nov 1971. Two incomplete colonies. 

85288: Sars Seamount, Drake Passage: 59°49'S, 
68°52'W, 512-622 m, Eltanin sta. 254, 10 Oct 1962. 
One colony lacking holdfast, and pieces of denuded 
axis (SEM 1647). 

85290: South of Falkland Islands, vicinity of 
Beauchene Is.: 53°08'S, 59°23'W, 567-578 m, Eltanin 
sta. 340, 3 Dec 1962. Two colonies, one attached to 
pebble. 

85298: Strait of Le Maire: 54°59'S, 64°53'W, 641- 
586 m, Eltanin sta. 970, 11 Feb 1964. Four small col- 
onies, somewhat damaged. (SEM 1648-1650.) 

85300: Tierra del Fuego, north of Cabo San Diego: 
53°32'S, 64°57'W, 124-119 m, Eltanin sta. 974, 12 
Feb 1964. 22 more or less damaged colonies. (SEM 
1926-1929.) 

85302: East edge of Burdwood Bank: 54°43'S, 
55°30'W, 1647-2044 m, Eltanin sta. 1592, 14 Mar 
1966. Two small colonies, attached to stones with al- 
cyoniids. (SEM 1652.) 

85306: Tierra del Fuego, off Cabo San Sebastian: 
53°06'S, 67°04'W, 86 m, RA^ Hero sta. 450, 5 Mar 
1970. Nine colonies, one attached to stone, one to peb- 
ble, others lacking holdfast. (SEM 1644, 1645, 1658- 
1660, 1921.) 

85308: Tierra del Fuego, off north entrance to Strait 
of Le Maire: 54°34'S, 64°10'W, 73 m, RA^ Hero sta. 
856, 20 Oct 1971. Five colonies, 2 with holdfast. 

85312: Tierra del Fuego, south of Isla de los Esta- 
dos: 54°55.6'S, 64°21.8'W, 303-358 m, RA/ Hero sta. 
893, 2 Nov 1971. Two colonies lacking holdfast. 

88359: Tierra del Fuego, east of Cabo San Sebas- 
tian: 53°15'S, 66°51'W, 79-80 m, Eltanin sta. 222, 27 
Sep 1962. One colony lacking holdfast. 

88362: South of Falkland Islands, vicinity of 
Beauchene Is.: 53°05'S, 59°31'W, 512-586 m, Eltanin 
sta. 339, 3 Dec 1962. Five colonies, three attached to 
pebbles. 

88747: East of Cape Horn: 56°06'S, 66°19'W, 384- 
494 m, Eltanin sta. 740, 18 Sep 1963. Two colonies, 
one lacking holdfast, one attached to pebble. 

88756: Tierra del Fuego, off north entrance to Strait 



* Eltanin station numbers 181-297 (Cruise 5, 1962) 
were duplicated in Cruise 21 (1965). 



VOLUME 109, NUMBER 1 



203 



of Le Maire: 54°34'S, 64°20'W, 91 m, RA' Hero sta. 
853, 20 Oct 1971. One colony lacking holdfast. 

88757: Tierra del Fuego, off north entrance to Strait 
of Le Maire: 54°29'S, 64°10'W, depth not recorded, YU 
V Hero sta. 855, 20 Oct 1971. One colony. 

88758: Tierra del Fuego, off north entrance to Strait 
of Le Maire: 54°33'S, 64°00'W, 84 m, RA' Hero sta. 
870, 24 Oct 1971. Two colonies lacking holdfast. 

88759: Tierra del Fuego, north of Isla de los Esta- 
dos: 54°39'S, 63°50'W, 135-137 m, RA^ Hero sta. 874, 
26 Oct 1971. Three colonies, two lacking holdfast. 

88760: Tierra del Fuego, off north entrance to Strait 



of Le Maire: 54°29'S, 63°50'W, 112 m, RA^ Hero sta. 
876, 27 Oct 1971. One colony lacking holdfast. 

88763: Tierra del Fuego, Strait of Le Maire: 
54°59.9'S, 64°50'W, 438-548 m, RA^ Hero sta. 895, 3 
Nov 1971. One colony lacking holdfast. 

89344: Between Tierra del Fuego and Falkland Is- 
lands: 52°35'S, 65°08'W, 128 m, Eltanin sta. 976, 13 
Feb 1964. Forty more or less complete colonies lack- 
ing holdfast. (SEM 1930-1934.) 

89345: Between Tierra del Fuego and Falkland Is- 
lands: 52°35'S, 65°08'W, 128 m, Eltanin sta. 976, 13 
Feb 1964. About 40 more or less completely denuded 
pieces of axis. 



The 123rd Annual Meeting of the Biological Society of Washington will be held 
on Tuesday, 7 May, 1996, at 12:00 noon in the Waldo Schmitt Room, National 
Museum of Natural History, Washington, D.C. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 

109(1):204. 1996 



INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 

Applications published in the Bulletin of Zoological Nomenclature 

The following Applications were published on 28 September 1995 in Vol. 52, 
Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on any of 
these applications is invited for publication in the Bulletin and should be sent to the 
Executive Secretary (I.C.Z.N.), % The Natural History Museum, Cromwell Road, 
London SW7 5BD, U.K. 

Case No. 

29 Al Patella longicosta Lamarck, 1819 (Mollusca, Gastropoda): proposed conser- 
vation of the specific name. 

2909 Glomeris Latreille, 1802 (Diplopoda): proposed conservation; Armadillo vul- 
garis Latreille, 1804 (Crustacea, Isopoda): proposed conservation of 
the specific name; and Armadillo Latreille, 1802 (Crustacea, Iso- 
poda): application for a ruling on its status. 

2894 Monstrilla Dana, 1849 and Thaumaleus Kr0yer, 1849 (Crustacea, Copepoda): 
proposed conservation. 

2967 Chaetodacus latifrons Hendel, 1915 (currently Bactrocera latifrons; Insecta, 
Diptera): proposed precedence of the specific name over that of 
Dacus parvulus Hendel, 1912. 

2933 Eudistoma Caullery, 1909 (Tunicata): proposed precedence over Paessleria 
Michaelsen, 1907. 

2966 Cyclodomorphus praealtus (Reptilia, Squamata): a proposal that availability 
of the specific name be taken from the intended description by Shea, 
1995. 



Opinions published in the Bulletin of Zoological Nomenclature 

The following Opinions were published on 28 September 1995 in Vol. 52, Part 3 
of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be ob- 
tained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History 
Museum, Cromwell Road, London SW7 5BD, U.K. 

Opinion No. 

1815. Chromadora Bastian, 1865 and Euchromadora de Man, 1886 (Nematoda): 

conserved by the designation of C. nudicapitata Bastian, 1865 as 
the type species of Chromadora. 

1816. Lithobius piceus L. Koch, 1862 (Chilopoda): specific name conserved. 

1817. Clavella Oken, 1815 and Pennella Oken, 1815 (Crustacea, Copepoda): con- 

served, and Pennella diodontis Oken, 1815: specific name con- 
served. 

1818. Rhopalosiphum monardae Davis, 1911 (currently Hyalomyzus monardae; 

Insecta, Homoptera): specific name conserved. 

1819. Bhatia Distant, 1908 (Insecta, Homoptera): Eutettix olivaceus Melichar, 1903 

confirmed as the type species. 

1820. A.A.H. Lichtenstein's (1796, 1797) Catalogus musei zoologici . . . Sectio 

Tertia Continens Insecta and D.H. Schneider's (1800) Verzeichniss 
einer Parthei Insekten . . . : suppressed, with conservation of some 
Lichtenstein (1796) names (Insecta and Arachnida). 

1821. Cliola (Hybopsis) topeka Gilbert, 1884 (currently Notropis topeka; Osteich- 

thyes, Cypriniformes): specific name conserved. 



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CONTENTS 

New species of mammals from northern South America: Bats of the genera Histiotus Gervais 

and Lasiurus Gray (Chiroptera: Vespertihonidae) Charles O. Handley, Jr. 1 

Identity and taxonomic status of the Mexican garter snake Thamnophis vicinus Smith, 1942 

(Reptilia: Serpentes: Natricidae) Douglas A. Rossman 10 

Review of the genus Cociella Whitley (Teleostei: Platycephalidae) with the description of three 

new species Leslie W. Knapp 17 

New locality records of freshwater decapods from Mexico (Crustacea: Atyidae, Cambaridae, 

and Palaemonidae) Gabino A. Rodriguez-Almaraz and Ernesto Campos 34 

Neocrangon zacae (Chace, 1937) synonymized with N. resima (Rathbun, 1902), and compared 

with TV. communis (Rathbun, 1 899) (Decapoda: Caridea: Crangonidae) Mary K. Wicksten 39 
Dalldorfia Rathbun, 1904 (Crustacea: Decapoda) from the Neogene of Japan 

Hiroaki Karasawa and Hisayoshi Kato 44 
Systematics and distribution of the genus Calocarides (Crustacea: Decapoda: Axiidae) 

Brian Kensley 53 
A new species of the axiid shrimp genus Acanthaxius from the Caribbean (Crustacea: Decap- 
oda: Thalassinidea) Brian Kensley 70 
A new genus for Anapagrides sensu De Saint Laurent-Dechance, 1966 (Decapoda: Anomura: 

Paguridae) and descriptions of four new species Patsy A. McLaughlin and Janet Haig 75 

Rediscovery of Cymodocella algonense from South Africa (Crustacea: Isopoda: Sphaeromati- 

dae) Brian Kensley and Mary Bursey 91 

Spelaeomysis villalobosi, a new species of mysidacean from northeastern Mexico (Crustacea: 

Mysidacea) Maria Elena Garcia-Garza, Gabino A. Rodriguez-Almaraz, 

and Thomas E. Bowman 97 
A new copepod species from California, U.S.A.: Hesperodiaptomus californiensis (Crustacea: 

Copepoda: Calanoida: Diaptomidae) Megan Scanlin and Janet W. Reid 103 

Orecturus amplus, a new species (Copepoda: Siphonostomatoida: Asterocheridae) from an 

alcyonacean in New Caledonia Arthur G. Humes 112 

Records of Enchytraeidae (Annelida: Oligochaeta) from west Florida. 1 . Mesenchytraeus, Cog- 

nettia, Bryodrilus, Hemienchytraeus, Henlea, and Buchholzia Brenda Healy 118 

Artacama valparaisiensis, a new species of Terebellidae (Annelida: Polychaeta) from subtidal 

soft bottoms of Valparaiso Bay, Chile Nicolas Rozbaczylo and Marco A. Mendez 138 

Review of Hennilepidonotus Uschakov, 1974, and two species of polynoid polychaetes (Lep- 

idonotinae) Marian H. Pettibone 143 

The Antarctic genus Callozostron and its relationship to Primnoella (Octocorallia: Gorgonacea: 

Primnoidae) Frederick M. Bayer 150 

International Commission on Zoological Nomenclature 204 

Notice: Annual Meeting 203 



6LH 
I 

.Vff 



PROCEEDINGS 
BIOLOGICAL SOCIETY 

OF 

WASHINGTON 






VOLUME 109 


NUMBER 2 


25 JUNE 1996 




ISSN 0006-324X 





THE BIOLOGICAL SOCIETY OF WASHINGTON 

1996-1997 
Ojficers 

President: Stephen D. Cairns Secretary: Carole C. Baldwin 

President-elect: Richard P. Vari Treasurer: T. Chad Walter 

Elected Council 

John Fornshell Rafael Lemaitre 

Alfred L. Gardner Diana Lipscomb 

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Custodian of Publications: Storrs L. Olson 



PROCEEDINGS 

Editor: C. Brian Robbins 

Associate Editors 

Classical Languages: George C. Steyskal Invertebrates: Jon L. Norenburg 

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):205-228. 1996. 

Three new species of precious coral (Anthozoa: Gorgonacea, 
genus Corallium) from Pacific waters 

Frederick M. Bayer 

Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — Two new species of Corallium from New Caledonia with the 
consolidated axial skeleton having smooth pits with beaded margins accom- 
modating the autozooids are described, Corallium thrinax with double-club 
sclerites, C. nix without. A third new species, C. kishinouyei, lacking smooth, 
well-defined axial pits and lacking double-club sclerites, is described from 
Cross Sea Mount south of Hawaii. Preliminary observations of axis formation 
are reported. 



Program BIOCAL conducted in 1985 by 
ORSTOM aboard RA^ Jean Charcot in the 
vicinity of New Caledonia obtained two 
species of the genus Corallium that cannot 
be assigned to any species heretofore de- 
scribed. Investigations of seamounts south 
of the Hawaiian Islands in 1993 by the 
Woods Hole Oceanographic Institution us- 
ing the research submersible Pisces V ob- 
tained another undescribed species. These 
are now placed on record. 

Suborder Scleraxonia Studer, 1887 

Family Coralliidae Lamouroux, 1812 

Genus Corallium Cuvier, 1798 

Synonymy. — See Bayer, 1956:70; 1964: 
466. 

Type species. — Madrepora rubra Lin- 
naeus, by subsequent monotypy, Lamarck, 
1801. 

Diagnosis. — Dimorphic Gorgonacea 
with continuous, solid, calcareous support- 
ing axis without hollow core. Coenenchy- 
mal sclerites including 6-, 7-, and 8-radiate 
capstans sometimes asymmetrically modi- 
fied as "opera-glass forms" or "double 
clubs," spindles, rods, and crosses. 

Remarks. — Following the conclusion by 
Lacaze-Duthiers (1864) that the calcareous 
skeleton of Corallium rubrum is composed 



of sclerites cemented inseparably to form a 
continuous, unsegmented axis, the genus 
Corallium has been classified in the sub- 
order Scleraxonia, in which the supporting 
axis unquestionably is composed of more 
or less completely fused sclerites as is the 
case in such families as the Paragorgiidae, 
Briareidae, and Melithaeidae. However, 
thin sections of the axis of Corallium ex- 
amined under the light microscope do not 
show convincing evidence of spicular struc- 
ture in spite of the statements to the con- 
trary by various authors up to the present 
time (Miiller 1910, Kiikenthal 1924, Hy- 
man 1940, Bayer 1956, Weinberg 1976). 

It has now been demonstrated that in 
Corallium rubrum formation of the axis at 
the apex of branches begins as an aggluti- 
nation of sclerites that become immersed in 
"rectilinear" calcite, which later comprises 
the predominant structural component 
(Grillo et al. 1993). This finding supports 
the inclusion of Corallium (family Coralli- 
idae) in the gorgonacean suborder Sclerax- 
onia. 

However, Lawniczak (1987) concluded 
that in C. johnsoni Gray the axis is com- 
posed entirely of fibrocrystalline calcium 
carbonate, without inclusion of sclerites. As 
it is illogical to assign species otherwise 
closely similar in morphological (dimorphic 



206 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



polyps) and skeletal (sclerites of virtually 
identical size and form) characters to dif- 
ferent orders (and, necessarily, different 
families) on the basis of a percieved differ- 
ence in skeleton formation observed in a 
limited sample of a rare and poorly known 
species, the family Coralliidae is retained in 
the Scleraxonia pending a wider investiga- 
tion of axis formation in as many species 
of Corallium as can be obtained for study. 
The description of Corallium thrinax, 
new species, was originally composed in 
French by Jeffrey Stefani, translated into 
English and edited by F. M. Bayer, so the 
species correctly bears joint authorship. The 
descriptions of C. nix and C. kishinouyei 
were prepared solely by the present author. 

Corallium thrinax Bayer & Stefani, 

new species 

Figs. 1-6 

Material examined. — New Caledonia, 
23°06.50'S, 167°53.74'E, BIOCAL 1985, 
R/V Jean Charcot station DW-50, 240 m, 
31-VIII-85. Sixteen colonies with holdfast, 
and some fragments: Holotype and 16 para- 
types (NMNH Paris); 8 paratypes, USNM 
96511 (SEM 1328-1331, 2416, 2466). 

Diagnosis. — Corallium branched dichot- 
omously and laterally with tendency to re- 
main in one plane, small, usually with no 
more than three or four bifurcations and a 
few short lateral twigs; autozooids forming 
low verrucae concentrated on one side of 
the branches; axis with distinct pits bor- 
dered by prominent beaded rim located be- 
neath autozooids. Predominant sclerites are 
6-radiate capstans and double clubs ("opera 
glasses") with smooth, globose heads. Col- 
onies white, sclerites colorless. 

Description. — The colonies are small, 
sparingly branched dichotomously, often 
with a few short lateral twigs (Fig. 1). They 
range from 3.7 to 6 cm in height, and be- 
tween 3 and 6 cm in width. The holdfast is 
a narrow basal expansion of the trunk. For 
the most part branching is in one plane, but 
in some colonies a branch may stand at 



nearly 90° from the plane of the principal 
fan. There are no anastomoses. In most 
cases there is a short main trunk, oval in 
cross section, about 0.6 cm X 1.2 cm in 
diameter, which bifuractes to produce prin- 
cipal branches approximately round in sec- 
tion, which in turn bifurcate as many as two 
or three times; a few short, blunt lateral 
twigs sometimes arise from one or more of 
the internodes between bifurcations. In 
some cases, principal branches 2-4 mm in 
diameter arise directly from the holdfast. 

The surface of the axis is longitudinally 
grooved (Fig. 2) and covered with minute 
tubercles ornamented with thorny projec- 
tions (Fig. 3 bottom). At the position of 
each autozooid, the axis has a distinct 
rounded pit 0.75 mm wide and 1.05 mm 
long (Fig. 3 top) bordered by a beaded rim 
that in some cases is interrupted by a nar- 
row gap for passage of a coelenteric canal 
(Fig. 2 bottom). 

Examination of the apex of the axis of a 
twig tip by SEM shows tubercles some re- 
sembling axial processes, others resembling 
the tubercles of sclerites (Fig. 4). Clearly 
recognizable sclerites partially embedded in 
axial calcite were not found on the single 
sample examined. This might be expected 
if that twig tip had ceased upward growth 
at time of collection. Owing to the destruc- 
tive nature of such preparations, no further 
samples have been made for the present 
purposes of taxonomic description. 

Autozooids form low, inconspicuous ver- 
rucae distributed predominantly on one face 
(the "front") of the colony, with only an 
occasional stray on the back surface (Fig. 
1). They are indistinct unless artificially 
stained with crystal violet. The tentacles are 
retracted flush with the coenenchyme, their 
bases forming an 8-lobed margin of the ver- 
rucal orifice. The autozooids are seated in 
distinct, rounded axial pits with a beaded 
rim (Figs. 2, 3). Siphonozooids are scarce 
and distributed randomly in the coenenchy- 
me, so inconspicuous that they can be dis- 
tinguished only by staining with crystal vi- 
olet. 



VOLUME 109, NUMBER 2 



207 




Fig. 1. Corallium thrinax, new species, a, Holotype colony; b, part of colony with zoanthid symbionts; c. 
Colony with autozooids stained with crystal violet, front surface; d. Colony with autOzooids stained with crystal 
violet, back surface; paratype colony USNM 965 1 1 . 



208 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 2. Corallium thrinax, new species. Apex of axis in front and lateral views SEM 1331, stereo pairs. 



The coenenchyme is so thin that the axial 
sculpture is exposed in areas subject to 
abrasion (Fig. 6 top). The longitudinal axial 
striations are clearly indicated through the 
coenenchyme, most clearly toward branch 



tips where the autozooids are closer togeth- 
er. 

Sclerites (Fig. 5) of the coenenchyme in- 
clude numerous 6-radiates up to 0.04 mm 
in length, many of which are modified as 



VOLUME 109, NUMBER 2 



209 




Fig. 3. Corallium thrinax, new species. Top, Axial pit at location of autozooid; Bottom, Axial protuberances. 
SEM 1331, stereo pairs. 



globose double clubs by asymmetrical hy- 
pertrophy of two of the rays to form a pair 
of smooth, spheroidal processes. At the sur- 
face of the coenenchyme these lie crowded, 
with the spheroidal processes directed out- 
ward (Fig. 6 bottom). Sclerites intermediate 



between double clubs and 6-radiates with 
the hypertrophied rays more or less strongly 
modified are not uncommon. Irregular 
forms occur but are extremely rare. No 
crosses or 8-radiates were observed, and 
only a single small 7-radiate was found in 



210 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




241701 10KV X500 60um 



Fig. 4. Corallium thrinax, new species. Top, Apex of axis showing axial protuberances and tubercles similar 
to those of sclerites. SEM 2416, stereo pair; Bottom, Surface of axis near apex showing axial protuberances and 
tubercles similar to those of sclerites. SEM 2417, stereo pair. 



VOLUME 109, NUMBER 2 



,211 




Fig. 5. Corallium thrinax, new species. Sclerites. SEM 1328-1330. 



3 preparations for SEM. Small rods and 6- the axis is white and the sclerites are col- 
radiates are present in the tentacles, the dis- orless. 

tal ones decreasing in size to about 0.03 Comparisons. — The colonial form of 

mm. Corallium thrinax is immediately distin- 

Colonies preserved in ethanol are white; guished from that of any other species of 



212 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 6. CoralUum thrinax, new species. Top, Tip of terminal twig; Bottom, Surface of coenenchyme showing 
predominance of double-club forms. SEM 2466, stero pairs. 



CoralUum known heretofore. Other nomi- 
nal species having autozooids seated in ax- 
ial pits with beaded rims are: C. stylaster- 
oides Ridley, 1882, from Mauritius; C. sal- 



omonense Thomson & Mackinnon, 1910, 
from Salomon in the Indian Ocean; and C. 
tortuosum Bayer, 1956, from Hawaii, which 
was reduced, perhaps prematurely, to a sub- 



VOLUME 109, NUMBER 2 



213 



species of salomonense (Bayer, 1993); C. 
inutile Kishinouye, 1903, from Japan; and 
C. nix new species, described herein. 

The axis of Corallium japonicum Kish- 
inouye also is marked by small pits beneath 
the autozooids (Kishinouye 1905:22), as is 
that of C elatius Ridley (Kishinouye, 1905: 
24) and C. rubrum (Linnaeus) (Grillo, 
Goldberg & Allemand, 1993:121, fig. Ic), 
but in those species (and probably others as 
well) the pits are not bordered by a distinct, 
beaded margin. Among those with beaded 
pits, C. stylasteroides, C. salomonense and 
C. tortuosum lack double clubs among the 
sclerites. C. inutile has double clubs of 
somewhat different outline and only 6-ra- 
diates but no 7- and 8-radiates, and differs 
further in its colonial form, pale pink axis 
and reddish cortex. C. borneense, which is 
salmon pink with red calices and quite dif- 
ferently branched, has double clubs of sim- 
ilar size and shape, but its autozooids form 
distinctly hemispherical verrucae and are 
not seated in rimmed pits in the axial skel- 
eton. 

Etymology. — thrinax from Greek Gpiva^, 
a three-pronged fork, in allusion to the 
forked braching of the colonies. Noun in 
apposition. 

Symbionts. — A few of the specimens are 
host to a balanoid barnacle that is partially 
overgrown by coral skeleton, and the 
branch tips of a few are overgrown with 
what appears to be a fasciculate hydroid. 
Near the base of some colonies there are 
symbiotic zoanthideans (Fig. lb), but these 
are not invariably present as seems to be 
the case in both C inutile and C. tortuosum. 

Corallium nix, new species 
Figs. 7-10 

Material examined. — New Caledonia, 
23°06.50'S, 167°53.74'E, BIOCAL 1985, 
RA^ Jean Charcot station DW-50, 240 m, 
31-VIII-85. One branch apparently part of 
a larger colony, holotype. Museum National 
d'Histoire Naturelle, Paris. 

Diagnosis. — Corallium irregularly branched; 



autozooids on all sides of the branches, 
seated in distinct pits in the axis, bordered 
by prominent beaded rim. Predominant 
sclerites are short 8-radiate capstans and 6- 
and 7-radiates some of which are weakly 
asymmetrical but in the form of double 
clubs; crosses extremely rare. Colonies 
white, sclerites colorless. 

Description. — As the only specimen ap- 
pears to be no more than a branch of a larg- 
er colony, it is not possible to provide a 
description of the general aspect of the 
complete colony. The single branch (Fig. 7) 
is sinuously curved, thick, approximately 
round in cross section, and 6.2 mm in great- 
est diameter at the base. Its shape suggests 
that branching probably proceeds roughly 
in one plane, as often is the case in species 
of Corallium. It produces short terminal 
branchlets mostly from two sides in roughly 
the plane of curvature, but a few small ones 
arise from the "front" of the branch. 

The surface of the axis is longitudinally 
grooved (Fig. 8), as usual covered with 
minute axial protuberances ornamented 
with thorny projections (Fig. 9). The axis 
has a distinct rounded pit 0.75 mm wide 
and 1.05 mm long at the position of each 
autozooid, bordered by a beaded rim that 
commonly is interrupted where a coelenter- 
ic canal passes through to connect with the 
gastrovascular cavity of the autozooid (Fig. 
8). Owing to the limitations of material 
available for study, an intact terminal twig 
was not prepared for examination by SEM 
to avoid damage to the unique type speci- 
men. As expected, no clearly recognizable 
sclerites partially embedded in the subapi- 
cal rectilinear calcite were observed. 

Autozooids are distributed on all sides of 
the branch and terminal branchlets (Fig. 
10). They retract flush within the rim of the 
axial pits, the bases of the infolded tentacles 
forming an 8-lobed margin of the closed 
verrucal apertures. Their distribution is 
most clearly revealed by staining with crys- 
tal violet, which stains the structures within 
the gastric cavities more darkly than the 
surrounding coenenchyme so that they con- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 7. Corallium nix, new species. Holotype branch with autozooids stained with crystal violet showing 
general distribution on front and back surfaces. Height overall, 5.75 cm. 



trast sharply with the paler cortex as the 
highly soluble stain dissipates (Fig. 7). 

The siphonozooids do not form distinct 
small verrucae and therefore are very dif- 
ficult to detect, but commonly one is locat- 
ed in the tapered groove extending from the 
autozooid, and scattered individuals can be 
seen in the coenenchyme between and 
around the autozooids. 

The coenenchyme is very thin, generally 
smooth but inconspicuously papillate in ar- 
eas protected from abrasion, suggesting that 
elsewhere the minute papillae may either 
have been rubbed off during collection, or 



have contracted completely as the result of 
contact with other objects in the trawl. In- 
conspicuous sinuous longitudinal grooves 
indicate the location of the principal coe- 
nenchymal canals. 

The predominant sclerites (Fig. 10) are 
short 8-radiate capstans reaching lengths of 
0.06-0.07 mm, none of which are elongated 
as belted spindles, and very short, wide 6- 
and 7-radiates 0.04-0.06 mm long, which 
may be weakly asymmetrical but not clear- 
ly modified as double clubs ("opera glass- 
es"); the shortest approach the tuberculate 
spheroid form. Crosses are absent or so 



VOLUME 109, NUMBER 2 



215 




Fig. 8. Corallium nix, new species. Top, Part of branch with autozooid pits; Bottom, Detail of autozooid 
pit. SEM 1334, stereo pairs. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




133422 10KV X1.50K'. £0.0um 133420 10KV )^1.50K 20.0um 



Fig. 9. Corallium nix, new species. Top, Surface of axis with axial protuberances; Bottom, Detail of axial 
protuberances. SEM 1334, stereo pairs. 



VOLUME 109, NUMBER 2 



217 




Fig. 10. Corallium nix, new species. Sclerites. SEM 1333. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



vanishingly rare as to escape notice in prep- 
arations made both for SEM and light mi- 
croscopy. The tentacles contain blunt rods 
derived from octoradiates, as shown by nu- 
merous intermediate forms. 

Skeleton and coenenchyme white, scle- 
rites colorless, translucent. 

Etymology. — Latin nix, nivis f., snow, in 
allusion to the white corallum. Noun in ap- 
position. 

Comparisons. — The distinct axial pits 
with beaded margins in which the auto- 
zooids of Corallium nix are recessed are 
similar to those of Corallium stylasteroides 
Ridley, C. salomonense (Thomson & 
Mackinnon), and C. thrinax Bayer & Ste- 
fani. However, the sclerites of C. stylaster- 
oides are smaller and not so sharply sculp- 
tured, and those of C. salomonense are both 
larger and more acutely sculptured. C. thri- 
nax differs conspicuously in its small, di- 
chotomously branched uniplanar growth 
form and its strongly asymmetrical double- 
club sclerites. 

Corallium kishinouyei, new species 
Figs. 11-19 

Material examined. — Cross Seamount: 
18°46.6'N, 158°14.8'W, 1145m. Pisces V 
dive PV235, coll. Scott France and Ewann 
Agenbroad, station CR105, 18 Aug 1993. 
Sample 1: One incomplete colony 11.5 cm 
tall with 2 detached branches, holotype 
USNM 94462 (SEM 2381, 2382, 2423). 
Sample 4: One incomplete colony consist- 
ing of one branch 1 2 cm tall and 4 smaller 
branches from the same colony, the largest 
11 cm tall, USNM 94463, paratype (SEM 
2285, 2455). 

Diagnosis. — Corallium sparsely branched, 
roughly in one plane; autozooids biserial 
and weakly directed toward one face of the 
colony, a few on front of branches, not seat- 
ed in distinct deep pits in the axis; axis lon- 
gitudinally grooved. Predominant sclerites 
are short 8-radiate capstans and short, stub- 
by 6- and 7 -radiates not modified as double 



clubs; crosses extremely rare. Colonies 
white in ethanol, sclerites colorless. 

Description. — Colonies sparingly 

branched in one plane, openly dichotomous 
or lateral (Fig. 11), the smaller terminal 
branchlets more or less clavate (Fig. 12). 
The stoutest main branch is nearly round, 
7.2 mm in diameter; the terminal branchlets 
are about 4 mm in diameter, with a tenden- 
cy to flattening in the plane of branching. 
Autozooids are situated biserially and di- 
rected slightly toward one face of the col- 
ony (the "front"), with occasional individ- 
uals also on the front face roughly between 
bilateral pairs, becoming more generally 
scattered on the stoutest branches. They re- 
tract to form low, moundlike verrucae about 
3 mm in diameter and at most 1 mm in 
height, the orifices with marginal lobes not 
necessarily as many as 8 but depending 
upon the degree of contraction. Numerous 
small, bluntly conical papillae less than 0.5 
mm in diameter, each with an apical pore, 
cover the surface of the coenenchyme 
around and between the autozooids, are in- 
terpreted as siphonozooids but not con- 
firmed by histological examination. 

The predominant coenenchymal sclerites 
(Fig. 13) are 8-radiate capstans from about 
0.05 mm up to 0.13 mm in length; 6-radi- 
ates are present also, and 7-radiates are un- 
common; crosses are present, as are a few 
irregular forms. The tentacles contain slen- 
der, bluntly pointed rods up to about 0.09 
mm in length, derived from the predomi- 
nant 8-radiate form. 

The mature axis is longitudinally striated 
but there are no deep, smooth pits beneath 
the autozooids. Depressions in the axis ac- 
commodating the autozooids are confined 
to apical regions, where they are irregular 
and do not have smooth bottoms and prom- 
inent beaded margins (Figs. 18, 19 top). As 
usual, the surface is ornamented with small, 
thorny protuberances (Fig. 16, bottom) sug- 
gesting the sculpture of the sclerites, but 
these are not indicative of sclerites embed- 
ded in the axial calcite. Locally, some of 
the axial protuberances are very prominent 



VOLUME 109, NUMBER 2 



219 




Fig. 11. Corallium kishinouyei, new species. Branches of holotype colony. 



(Fig. 17, bottom) and resemble certain scle- 
rites of other scleraxonians that do not oc- 
cur in Corallium (e.g., the medullar scle- 
rites of Paragorgid). 



Owing to the limitations of the speci- 
mens available for study, only three apical 
samples from the axis from terminal twigs 
were prepared for examination by SEM, 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 12. Corallium kishinouyei, new species. Tip of terminal twig with coenenchyme intact. SEM 2382, 
stereo pair. 



one from USNM 94463 (Figs. 18, 19) and 
two from USNM 94462 (Figs. 14-16). Of 
the latter, the distal tip of the axis is in one 
case flattened and blade-like, the two flat 
faces each with two narrow ridges (Fig. 14, 
bottom) that probably accommodated the 
major longitudinal coenenchymal canals; 
in the other sample it is tapered, bluntly 
pointed, flattened but without the promi- 
nent longitudinal ridges (Fig. 15). The one 
axis tip examined from USNM 94463 is 
tapered, bluntly pointed, roughly prismat- 



ic, flattened on the "back" side, with ir- 
regular longitudinal ridges and intervening 
rows of irregular pits (Figs. 18, 19 top), 
the larger of which doubtless were situated 
beneath autozooids; the smaller pits are too 
small and too close together to underlie au- 
tozooids. 

In all cases, the apex of the axis is com- 
posed largely of sclerites bound together by 
intervening non-spicular calcite (Figs. 16 
top, 17 top, 19 bottom). This agrees with 
the conclusion of Grillo et al. (1993:127) 



VOLUME 109, NUMBER 2 



221 




Fig. 13. Corallium kishinouyei, new species. Sclerites. SEM 2285. 



that in C. rubrum "sclerites are incorporat- 
ed to form the core of the nascent skele- 
ton," subsequently becoming overlain by 
non-spicular calcite as the girth of the axis 
increases. 

The color of the colonies in ethanol ini- 



tially was distinctly yellowish white but the 
yellowish tint soon faded in alcohol; the 
sclerites are colorless. Those of the larger 
branches are predominantly opaque white, 
those of the terminal branches glass-clear, 
but there is no perceptible difference in 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



J"- i^v '? 



<^4hM 









24S331 10KV Xi00' ' .30mm I 242333 10KV X100 .30mm 




Fig. 14. Corallium kishinouyei, new species, axis of USNM 94462, preparation \. Top, Apex; Bottom, Distal 
portion just below apex. SEM 2423, stereo pairs. 



VOLUME 109, NUMBER 2 



223 




Fig. 15. Corallium kishinouyei, new species, axis of USNM 94462, preparation 2. Top, Apex from side; 
Bottom, Apex from front. SEM 2480, stereo pairs. 



224 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 16. Corallium kishinouyei, new species, axis of USNM 94462, preparation L Top, Sclerites embedded 
in non-spicular calcite at apex, SEM 2480; Bottom, Axial protuberances, SEM 2423. Stereo pairs. 



VOLUME 109, NUMBER 2 



225 




Fig. 17. Corallium kishinouyei, new species, axis of USNM 94462, preparation 1. Top, Surface of axis near 
apex, showing embedded sclerites and axial protuberance; Bottom, Surface of axis with typical axial protuber- 
ances and unusually large spiny projections. SEM 2480, stereo pairs. 



226 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 18. Corallium kishinouyei, new species, axis of USNM 94463. Top, Apex of axis, side view; Bottom, 
Apex of axis, front view. SEM 2455, stereo pairs. 



VOLUME 109, NUMBER 2 



227 




Fig. 19. Corallium kishinouyei, new species, axis of USNM 94463. Top, surface showing pits without beaded 
rim; Bottom, Surface near apex, showing embedded sclerites. SEM 2455, stereo pairs. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



crystal structure that might account for this 
difference in opacity. 

Etymology. — Named in honor of Kama- 
kichi Kishinouye, pioneer Japanese inves- 
tigator of precious corals. 

Comparisons. — The openly branched, 
uniplanar colonial form of Corallium kish- 
inoueyi is unlike any other species de- 
scribed so far, and the sclerites reach a larg- 
er size than in any species heretofore 
known. 

Acknowledgments 

I thank Dr. Bertrand Richer de Forges 
and Dr. Alain Crosnier of the Museum Na- 
tional d'Histoire Naturelle, Paris, for the 
opportunity to report upon octocorals col- 
lected around New Caledonia under auspi- 
ces of ORSTOM. Dr. Scott France, then of 
the Woods Hole Oceanographic Institution, 
provided the specimens from Cross Sea- 
mount. I am grateful to Jeffrey Stefani for 
the description of Corallium thrinax, pre- 
pared during the period when he assisted 
me in studies of octocorals from New Cal- 
edonia. The scanning electron micrographs 
were made by Walter R. Brown, head of the 
SEM laboratory of the National Museum of 
Natural History. Molly K. Ryan assembled 
and mounted the plates of isolated sclerites. 

Literature Cited 

Bayer, F. M. 1956. Descriptions and redescriptions of 
the Hawaiian octocorals collected by the U.S. 
Fish Commission steamer "Albatross" (2. Gor- 
gonacea: Scleraxonia). — Pacific Science 10:67- 
95. 

. 1964. The genus Corallium (Gorgonacea: 

Scleraxonia) in the western North Atlantic 



Ocean. — Bulletin of Marine Science of the Gulf 
and Caribbean 14:465-478. 

. 1993. Generic reassignment and affinities of 

Sympodium salomonense Thomson & Mac- 
kinnon (Coelenterata: Octocorallia). — Precious 
corals and octocoral research 1:14-19. 

Grillo, M.-C, W. M. Goldberg, & D. Allemand. 1993. 
Skeleton and sclerite formation in the precious 
red coral Corallium rubrum. — Marine Biology 
117:119-128. 

Hyman. L. H. 1940. Pp. 1-716 in The invertebrates. 
Protozoa through Ctenophora. McGraw Hill, 
New York. 

Kishinouye, K. 1903. Preliminary note on the Coral- 
liidae of Japan. — Zoologische Anzeiger 26: 
623-626. 

. 1905. Notes on the natural history of cor- 
als. — Journal of the Imperial Fisheries Bureau 
14:1-32. 

Kiikenthal, W. 1924. Gorgonaria. — Das Tierreich 47: 
1-478. 

Lacaze-Duthiers, H. 1864. Histoire naturelle du corail. 
J. B. Balliere et Fils, Paris, pp. 1-371. 

Lamarck, J. B. P. A. 1801. Systeme des animaux sans 
vertebres, ou tableau general des classes, des 
ordres et des genres de ces animaux. Paris, chez 
I'auteur, pp. 1-432 + 402bis. 

Lawniczak, A. 1987. Les modalites de croissance de 
I'axe calcaire chez Corallium johnsoni. (Cni- 
daria: Gorgonaria: Scleraxonia). — Senckenber- 
giana maritima 19:149—161. 

Miiller, R. 1910. Uber die Bildung des Achsenskelet- 
tes von Corallium. — Mittheilungen aus der 
Zoologischen Station zu Neapel zugleich ein 
Repertorium fiir Mittelmeerkunde 20:101-107. 

Ridley, S. O. 1882. On the arrangement of the Cor- 
alliidae, with descriptions of new or rare spe- 
cies. — Proceedings of the Zoological Society of 
London 1882:221-233. 

Thomson, J. A., & D. L. Mackinnon. 1910. Alcyon- 
arians collected on the Percy Sladen Trust Ex- 
pedition by Mr. J. Stanley Gardiner. Part 2. The 
Stolonifera, Alcyonacea, Pseudaxonia, and Ste- 
lechotokea. — Transactions of the Linnean So- 
ciety of London (2) 13(2): 165-211. 

Weinberg, S. 1976. Revision of the common Octo- 
corallia of the Mediterranean circalittoral. I. 
Gorgonacea.— Beaufortia24(313):63-104. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):229-235. 1996. 

Reconsiderations of the species status of some South American 
Planarians (Platyhelminthes: Tricladida: Paludicola) 

Ronald Sluys 

Institute for Systematics and Population Biology, Zoological Museum, 

University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, 

The Netherlands & UNISINOS, Sao Leopoldo, Brazil 

Abstract. — Reexamination of the type material of Girardia veneranda (Mar- 
tins, 1970) revealed that this nominal species is very similar to G. chilla (Mar- 
cus, 1954). Therefore, G. veneranda is determined to be a junior synonym of 
G. chilla. For the first time a morphological description is provided of speci- 
mens collected in 1970 near Buenos Aires and identified as G. anceps (Kenk, 
1930). The features of these G. anceps specimens conform in many details 
with Bohmig's (1902) redescription of this species. The material examined is 
compared also with more recent descriptions, from other localities, of G. anceps 
and with taxonomically related species. 



Adequate assessment of the world's bio- 
diversity depends on up to date knowledge 
on the diversity of species, their character- 
istics, and their distribution. The literature 
often features species descriptions and iden- 
tifications that are poorly documented or 
cannot stand scrutiny. In each of these cir- 
cumstances published results need to be 
checked on available, original material. 
This paper analyzes and discusses the spe- 
cies status of several South American pla- 
narians from the genus Girardia Ball, 1974. 
Planarians exemplify the situation that spe- 
cies are frequently characterized by a 
unique combination of diagnostic features 
instead of showing apomorphic characters 
(see Sluys 1991 for a review on species 
concepts). The variability that is encoun- 
tered in these softbodied animals has made 
species recognition in Girardia often very 
complex and difficult. 

De Vries & Sluys (1991) analyzed the 
phylogenetic relationships between the var- 
ious genera and subgenera within the Du- 
gesiidae Ball, 1974. They showed that the 
subgenera DMge5/a Girard, 1850, Schmidtea 
Ball, 1974, and Girardia Ball, 1974, are 
separate phylogenetic groups and that Du- 



gesia is not closely related to the other two 
subgenera. Therefore, they proposed to 
raise each of these subgenera to the rank of 
genus. The postulated apomorphies for the 
genus Girardia are a pigmented pharynx 
and a high triangular head shape with point- 
ed auricles, being features that characterize 
many paludicolans from North America, 
Central and South America, and the Carib- 
bean. 

The present paper for the first time pro- 
vides morphological evidence supporting 
the suggestions of Ball (1974) and Kenk 
(1974) that G. veneranda (Martins, 1970) 
from the State of Sao Paulo might be iden- 
tical with G. chilla (Marcus, 1954) from 
southern Chile. 

In 1970 Duran-Troise & De Lustig pub- 
lished the chromosome portrait of Girardia 
anceps (Kenk, 1930) from Buenos Aires, 
Argentina. Unfortunately, these authors did 
not provide a taxonomic description sup- 
porting the identification of their material. 
Apart from the fact that the identification of 
Duran-Troise & De Lustig needs proper 
documentation, examination of their mate- 
rial is also interesting in the light of Mor- 
etto's (1991) remark that G. anceps does 



230 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

cb 




100pm 



Figs. 1-2. Girardia chilla. 1, MZUSP 5829-5830. Sagittal reconstruction of the copulatory apparatus, be: 
bursal canal; cb: copulatory bursa; gl: gland; od: oviduct; pp: penis papilla; sg: shell gland; sv: seminal vesicle; 
vd: vas deferens; 2, MZUSP 5819-5822. Sagittal reconstruction of the copulatory apparatus, be: bursal canal; 
cb: copulatory bursa; gl: gland; od: oviduct; pp: penis papilla; sg: shell gland; vd: vas deferens. 



not occur near Buenos Aires. Recently, 
Puccinelli & Deri (1991) also described the 
chromosomes of G. anceps from Buenos 
Aires, unfortunately without an anatomical 
description of their specimens or mention- 
ing the deposition of voucher specimens. 

Taxonomic Section 

Girardia veneranda (Martins, 1970) 
Figs. 1, 2 

Material examined. — Although the major 
part of the material studied by Martins 
(1970) was examined, the present re-de- 
scription is based mainly on the following 
preparations [housed in the Museu de Zool- 
ogia, Universidade de Sao Paulo, Se^ao 
Helmintologia (MZUSP)]: 5813-5816 (sag- 
ittal sections on 4 slides; slide no. 5813 
missing), 5817-5818 (sagittal sections on 2 
slides), 5819-5822 (sagittal sections on 4 
slides), 5823-5825 (transverse sections on 
3 slides), 5826-5828 (sagittal sections on 3 
slides), 5829-5830 (sagittal sections on 2 
slides), 5831-5834 (horizontal and trans- 
verse sections on 4 slides). 



Description. — In the sectioned material 
no pigmentation appears to be present un- 
derneath the pharynx epithelium. The 
mouth opening is at about one-third of the 
distance between the hind wall of the pha- 
ryngeal pocket and the root of the pharynx. 

The testes are principally situated dorsal- 
ly, but large testes in the posterior end of 
the body may occupy the entire dorso-ven- 
tral space; the follicles extend from directly 
posterior to the ovaries to almost the pos- 
terior body end. The paired ovaries lie at a 
short distance behind the brain. 

The vasa deferentia, which form well de- 
veloped false seminal vesicles, recurve at 
about the level of the gonopore and open 
separately into the antero-lateral portions of 
the intrapenial seminal vesicle. This spher- 
ical or irregularly shaped seminal vesicle 
occupies most of the penis bulb and papilla. 
The wall of the vesicle is lined with a nu- 
cleate epithelium that is pierced by numer- 
ous openings of highly abundant cyano- 
philous glands, which lie within the penis 
bulb as well as directly outside of the bulb. 



VOLUME 109, NUMBER 2 

sg od 



231 




eg eg 



od sg 




Figs. 3—4. Girardia anceps. 3, ZMA V. PI. 871.5. Sagittal reconstruction of the copulatory apparatus, be: 
bursal canal; cb: copulatory bursa; eg: cyanophilie glands; ed: ejaeulatory duet; eg: erythrophilic glands; od: 
oviduct; pp: penis papilla; sg: shell gland; vd: vas deferens; 4, ZMA V. PL 871.1. Sagittal reconstruction of the 
copulatory apparatus, be: bursal canal; cb: copulatory bursa; eg: cyanophilie glands; ed: ejaeulatory duct; eg: 
erythrophilic glands; od: oviduct; pp: penis papilla; sg: shell gland; vd: vas deferens. 



The secretion is also abundantly present in 
the seminal vesicle. 

The penis papilla is a stubby cone, cov- 
ered with a nucleate epithelium which is 
underlain with a layer of circular and lon- 
gitudinal muscle, successively. 

The copulatory bursa is situated directly 
anterior to the penis bulb. From the bursa, 
the bursal canal curves smoothly towards 
the postero-dorsal portion of the atrium; the 
canal is lined with cuboidal cells bearing 
long cilia. The canal is surrounded by a lay- 
er of intermingled circular and longitudinal 
muscle. The two oviducts fuse to form a 
very short common oviduct that immediate- 
ly communicates with the bursal canal. 
Ventral to the opening of the common ovi- 
duct the bursal canal receives the erythro- 
philous secretion of shell glands. Another 



set of eosinophilous glands discharges their 
secretion into the gonopore. 

Girardia anceps (Kenk, 1930) = Planaria 

dubia Borelli, 1895 

Figs. 3, 4 

Material examined. — The material ex- 
amined consists of the material collected 
and described by Duran-Troise & De Lustig 
(1970) but identified by Prof. Benazzi. This 
material consists of the following series of 
slides, now housed in the Zoological Mu- 
seum, Amsterdam (ZMA): V. PI. 871.1: 
sagittal sections of one animal on 3 slides; 
V. PI. 871.2: sagittal sections on 2 slides; 
V. PL 871.3: sagittal sections on 2 slides; 
V. PI. 871.4: sagittal sections on 2 slides; 
V. PI. 871.5: sagittal sections on 4 slides; 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



V. PI. 871 .6: transverse sections on 4 slides; 
V. PI. 871.7: horizontal sections on 3 slides; 
V. PL 871.8: sagittal sections on 1 slide. 

Description. — The slides revealed that 
the animals are pigmented on dorsal and 
ventral body surface. The sections also un- 
equivocally showed the pharynx to be pig- 
mented. The mouth opening is located at 
the hind end of the pharyngeal pocket. 

The numerous small testes are situated 
ventrally, from directly behind the ovaries 
extending to almost the posterior body end. 
The ovaries are located at some distance be- 
hind the brain. 

The vasa deferentia recurve at the level 
of the penis bulb, penetrating it from the 
lateral sides. Within the bulb the ducts ex- 
pand in diameter and within the penis pa- 
pilla they fuse to form a broad ejaculatory 
duct. The intrapenial parts of the vasa de- 
ferentia and the ejaculatory duct are lined 
with a well developed, nucleate epithelium 
and are surrounded by a thin layer of cir- 
cular muscle. The penis papilla is lined with 
a flat, infranucleate epithelium. Ejaculatory 
duct and penis epithelium are pierced by 
ducts of numerous eosinophilic glands, 
while cyanophilous glands open into the in- 
trabulbar seminal vesicles. 

The narrow male atrium communicates 
with the common atrium; there is no dis- 
tinct female atrium. The atria are lined with 
an infranucleate epithelium. Especially the 
posterior wall of the common atrium is 
pierced by openings of numerous eosino- 
philic glands; the glandular elements being 
broadly distributed in the surrounding pa- 
renchyma. 

The bursal canal is more or less "an- 
gled"; the short section of the canal that 
sharply turns towards the dorsal part of the 
atrium receives the openings of erythro- 
philic shell glands. The bursal canal is lined 
with an infranucleate, ciliated epithelium. 
The musculature of the bursal canal either 
consists predominantly of circular muscles 
(as in preparations V. PL 871.1 and 871.5) 
or contains intermingled circular and lon- 
gitudinal fibres (as in preparation V. PL 



871.3). The infranucleate oviducts join to 
form a very short common oviduct that 
opens into the rear wall of the bursal canal, 
just dorsally to the angled section of the 
canal. 

In specimen V.Pl.871.1 remnants of a 
spermatophore project from the ejaculatory 
duct through the gonopore. 

Discussion 

The above description of G. veneranda 
differs in some important details from that 
given by Martins (1970). Although she de- 
scribed the vasa deferentia as opening "... 
into a single and spacious true seminal ves- 
icle . . . ," the illustration of the copulatory 
apparatus (Martins 1970, fig. 5) seems to 
indicate that the ducts open into a sort of 
intrapenial papilla. However, this sugges- 
tion of an intrapenial papilla merely results 
from an invagination or fold of the irregu- 
larly shaped seminal vesicle. In reality the 
vasa deferentia penetrate separately the lat- 
eral wall of the vesicle, as correctly depict- 
ed in figs. 6 and 7 of Martins. 

According to Martins, the oviducts open 
separately into the bursal canal. However, 
detailed study of both sagittal and trans- 
verse sections revealed that the oviducts 
fuse to an extremely short common duct. 
This situation is already indicated in Mar- 
tin's fig. 8 but was interpreted differently in 
her fig. 5 and in the text description of the 
copulatory apparatus. 

The present re-analysis of the type ma- 
terial of G. veneranda reveals that the mor- 
phology of the reproductive apparatus is 
strikingly similar to that of G. chilla, as de- 
scribed by Marcus (1954) and Hyman 
(1959). Both species show the spacious in- 
trapenial seminal vesicle, receiving the 
abundant cyanophilous secretion. Accord- 
ing to Hyman (1959) eosinophilous glands 
open into the vesicle, but this does not 
agree with my observations and those of 
Marcus (1954). The only difference to be 
noted between the G. veneranda specimens 
and G. chilla is that the latter was described 



VOLUME 109, NUMBER 2 



233 



with numerous glands opening through the 
entire epithehum of the common atrium. In 
the G. veneranda material the glands were 
not as abundantly present as described by 
Marcus or Hyman. According to Marcus 
these glands are cyanophilic but Hyman de- 
scribed them as eosinophilic, which con- 
forms to my observations. 

In view of the above, it is here concluded 
that G. veneranda is essentially similar to 
G. chilla and that therefore G. veneranda 
must be considered a junior synonym of G. 
chilla. 

Because Borelli's (1895) description of 
Planaria dubia [=G. anceps] is rather su- 
perficial and therefore does not allow de- 
tailed comparison with the material de- 
scribed in the present paper, it is here ac- 
cepted that Bohmig's (1902) more compre- 
hensive description applies to Borelli's 
species. This axiom is open for criticism, of 
course, since Borelli's material came from 
Asuncion, Paraguay and Bohmig's speci- 
mens were collected near Buenos Aires, Ar- 
gentina (but it must be noted that Bohmig 
did examine Borelli's preparations). How- 
ever, the main purpose of the present dis- 
cussion is to compare my material with (1) 
Bohmig's account of G. anceps, (2) more 
recent descriptions of the species, and (3) 
descriptions of taxonomically related spe- 
cies. 

Bohmig mentioned for G. anceps the fol- 
lowing characteristics: (1) recurved vasa 
deferentia, (2) infranucleate epithelium lin- 
ing the atrium and the penis papilla, (3) cir- 
cular muscles around ejaculatory duct and 
intrabulbar parts of the vasa deferentia, (4) 
cyanophilous glands discharging into these 
intrabulbar seminal vesicles, (5) ejaculatory 
duct receiving the secretion of eosinophi- 
lous glands, (6) proximal, anterior section 
of bursal canal funnel-shaped, (7) posterior 
two-thirds of bursal canal lined with infra- 
nucleate epithelium, (8) oviducts opening 
separately into bursal canal, (9) large num- 
ber of eosinophilous glands (different from 
the shell glands) discharging into the atri- 
um, (10) ventral testes, (11) dorsal surface 



dark brown with a pale mid-dorsal stripe. 
Bohmig's diagrammatic reconstruction of 
the copulatory apparatus of his G. anceps 
specimens has been redrawn by Cazzaniga 
& Curino (1987, fig. 2). 

It is evident that the material of Duran- 
Troise & De Lustig (1970), as described in 
the present paper, conforms in many details 
with Bohmig's account of G. anceps, the 
only possible difference being the openings 
of the oviducts into the bursal canal. With 
respect to the openings of the oviducts, 
however, it must be noted that Bohmig's re- 
construction (cf. Cazzaniga & Curino 1987, 
fig. 2) suggests oviducts opening very 
closely together into the bursal canal. Fur- 
thermore, it is important to note that in one 
particular Girardia species some specimens 
may have oviducts opening separately (but 
closely together) into the bursal canal, 
whereas in other animals the oviducts fuse 
to form a very short common oviduct. The 
same phenomenon was mentioned by Caz- 
zaniga & Curino for their specimens of G. 
anceps. 

Especially striking similarities between 
our material and Bohmig's account are the 
infranucleate epithelia of penis papilla, atri- 
um, and bursal canal. It is therefore dis- 
turbing that Bohmig found Borelli's mate- 
rial with nucleate linings of penis papilla 
and atria (this may cast doubt on the ident- 
icality of Bohmig's and Borelli's material; 
neither does Borelli depict the funnel- 
shaped section of the bursal canal where it 
communicates with the bursa). Cazzaniga 
& Curino also found in their specimens a 
bursal canal lined with a nucleate epitheli- 
um, but the atrium and penis papilla in 
these animals were provided with an in- 
franucleate lining epithelium. These differ- 
ences in nucleate/infranucleate epithelia, as 
reported by various authors, may simply re- 
flect different states of development of the 
copulatory apparatus, only fully mature in- 
dividuals with all their epithelia infranu- 
cleate. In Dugesia gonocephala (Duges, 
1830) De Vries (1984) found that nucle- 



234 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



ation of the bursal canal depends on the 
state of development. 

Apart from the differences mentioned 
above, the material of Cazzaniga & Curino 
is essentially similar to that studied by me. 
They mention also the infranucleate epithe- 
lium of the oviducts, which are surrounded 
by successively a layer of circular muscle 
and nuclei. In contrast to my material, Caz- 
zaniga & Curino reported the pharynx to be 
unpigmented. 

In view of the above, it is concluded that 
the specimens from Buenos Aires analyzed 
by Duran-Troise & De Lustig correspond 
with Bohmig's account of G. anceps from 
the same locality and also with the material 
described by Cazzaniga & Curino from Ba- 
hia Blanca. This contrasts with Moretto's 
(1991) conclusion that G. anceps has been 
recorded incorrectly from Buenos Aires. 

The specimens that Kawakatsu & Rova- 
sio (1992) used for their redescription of G. 
anceps differs in the following features 
from my material, Bohmig's account, and 
the description by Cazzaniga & Curino: (1) 
dorsal surface uniform blackish brown, 
without light mid-dorsal stripe, (2) pharynx 
unpigmented, (3) mouth opening somewhat 
anterior of posterior end of pharyngeal 
pocket (as suggested by their fig. 3B), (4) 
divided atrium, (5) male and common atri- 
um lined with nucleate epithelium, (6) bur- 
sal canal with nucleate epithelium, (7) bur- 
sal canal provided with very thick muscle 
coat. Each of these characters is known to 
vary between specimens of a single species 
and it is debatable whether a combination 
of these features indicates a species that is 
different from G. anceps and possibly from 
any other known species of Girardia. 

Several authors have discussed the simi- 
larities of G. anceps with G. sanchezi (Hy- 
man, 1959) and G. festae (BoreUi, 1898) 
and the species status of these taxa (Hyman 
1959, Cazzaniga & Curino 1987, Moretto 
1991, Kawakatsu & Rovasio 1992) [recent- 
ly, Curino & Cazzaniga (1993) argued that 
the original spelling G. festae was correct 
and that later emendations to G. festai were 



invalid]. Sluys (1992) considered G. san- 
chezi to be a junior synonym of G. festae 
because both were described with abundant 
glands opening into the common atrium, 
unicellular glands opening into the bursal 
canal, and with penial glands discharging 
through the epithelium of the penis papilla. 
Moreover, G. sanchezi and G. festae were 
both described with a pale, longitudinal 
mid-dorsal stripe. However, the present 
study shows that several of these features 
occur also in G. anceps, suggesting that G. 
festae might in turn be a junior synonym of 
G. anceps. Although G. anceps and G. fes- 
tae are similar in several aspects of their 
reproductive system, G. festae is different 
in that it has unicellular glands piercing the 
entire length of the nucleate bursal canal, 
and nucleate linings of penis papilla and 
atria. 

Thus, the conclusion of the present study 
is that G. sanchezi is a junior synonym of 
of G. festae and that G. festae and G. an- 
ceps are valid species. This conclusion is 
different from that reached by Cazzaniga & 
Curino (1987) and Kawakatsu & Rovasio 
(1992) that G. sanchezi is a valid taxon. 
However, the conclusion of these workers 
was based on a comparison of G. sanchezi 
with G. anceps, whereas the present study 
suggests that G. sanchezi is morphological- 
ly closer to G. festae. There is one feature 
that argues against G. sanchezi being a ju- 
nior synonym of G. festae, thus illustrating 
the point made in the introduction that spe- 
cies recognition in Girardia can be very 
complex: G. festae has recurved vasa de- 
ferentia (cf. Sluys 1992, fig. 2) while this is 
not the case in G. sanchezi (cf. Hyman 
1959, fig. 5). 

Acknowledgments 

Prof. Dr. M. Benazzi (Pisa, Italy) is 
thanked for making available the material 
of G. anceps described in this paper and for 
depositing the preparations in the Zoologi- 
cal Museum, Amsterdam. Prof. Dr. E. M. 
Froehlich (Sao Paulo, Brazil) is thanked for 



VOLUME 109, NUMBER 2 



235 



arranging the loan of the type material of 
G. veneranda. Part of this study was pre- 
pared while I stayed as a Visiting Professor 
at the Universidade do Vale do Rio dos Si- 
nos, Sao Leopoldo, Brazil; UNISINOS and 
the Brazilian Council for Scientific and 
Technological Development (CNPq) are 
thanked for making this visit possible. The 
research of the author has been made pos- 
sible, in part, by a grant from the Nether- 
lands Organization for Scientific Research- 
NWO (Biodiversity in Disturbed Ecosys- 
tems Programme). 

Literature Cited 

Ball, I. R. 1974. A contribution to the phylogeny and 
biogeography of the freshwater triclads (Platy- 
helminthes, Turbellaria). Pp. 339-401 in N. W. 
Riser & M. P. Morse, eds.. Biology of the Tur- 
bellaria, McGraw-Hill, New York, 530 pp. 

Bohmig, L. 1902. Turbellarien: Rhabdocoeliden und 
Tricladen. Hamburger Magalhaenische Sam- 
melreise, L. Friederichsen & Co., Hamburg, 30 
pp. + pis. I-II. 

Borelli, A. 1895. Viaggio del dott. Alfredo Borelli 
nella Repubblica Argentina e nel Paraguay XIII. 
Planarie d'acqua dolce. — Bollettino dei Musei 
di Zoologia ed Anatomia comparata della R. 
Universita di Torino 10:1-6. 

Cazzaniga, N. J., & A. C. Curino. 1987. On Dugesia 
anceps (Kenk, 1930) from Argentina (Turbel- 
laria: Tricladida).— Bollettino di Zoologia 54: 
141-146. 

Curino, A. C, & N. J. Cazzaniga. 1993. A new spe- 
cies of freshwater planarian from Chile (Platy- 
helminthes: Tricladida), with a nomenclatural 
note on Girardia festae (Borelli, 1898). — Pro- 
ceedings of the Biological Society of Washing- 
ton 106:633-644. 

De Vries, E. J. 1984. On the species of the Dugesia 
gonocephala group (Platyhelminthes, Turbellar- 



ia, Tricladida) from Greece. — Bijdragen tot de 
Dierkunde 54:101-126. 

, & R. Sluys. 1991. Phylogenetic relationships 

of the genus Dugesia (Platyhelminthes, Triclad- 
ida, Paludicola). — Journal of Zoology, London 
223:103-116. 

Duran-Troise, G., & E. S. De Lustig. 1970. Cariologia 
della planaria Dugesia anceps. — Caryologia 23: 
455-459. 

Hyman, L. H. 1959. On two fresh-water planarians 
from Chile. — American Museum Novitates 
1932:1-11. 

Kawakatsu, M., & R. A. Rovasio. 1992. Redescrip- 
tion of Dugesia anceps (Kenk, 1930) (Turbel- 
laria: Tricladida: Paludicola), from the vicinity 
of Cordoba, Argentina. — Proceedings of the 
Japanese Society of Systematic Zoology 48:7- 
23. 

Kenk, R. 1974. Index of the genera and species of the 
freshwater triclads (Turbellaria) of the world. — 
Smithsonian Contributions to Zoology 183:1- 
90. 

Marcus, E. 1954. Reports of the Lund University 
Chile Expedition 1948-49. 11. Turbellaria. 
Lunds Universitets Arsskrift N. F. Avd. 2, Bd 
49, Nr. 13:1-114. 

Martins, M. E. Q. P. 1970. Two new species of Du- 
gesia (Tricladida Paludicola) from the State of 
Sao Paulo, Brazil. — Anais da Academia Brasi- 
leira de Ciencias 42:113-118. 

Moretto, H. J. A. 1991. On the taxonomic status of a 
Dugesia species from Buenos Aires (Platyhel- 
minthes: Turbellaria: Tricladida). — Hydrobiolo- 
gia 227:353-354. 

Puccinelli, I., & P. Deri. 1991. Comparative karyolog- 
ical analysis of some American planarians be- 
longing to the genus Dugesia (subgenus Gir- 
ardia) (Platyhelminthes, Tricladida). — Caryol- 
ogia 44:225-232. 

Sluys, R. 1991. Species concepts, process analysis, 
and the hierarchy of nature. — Experientia 47: 
1162-1170. 

. 1992. Synopsis of the freshwater triclads of 

the Caribbean (Platyhelminthes, Tricladida, Pal- 
udicola). — Studies on the Natural History of the 
Caribbean Region 71:1-23. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 

109(2):236-247. 1996. 

Pycnophyes parasanjuanensis, a new kinorhynch 

(Kinorhyncha: Homalorhagida: Pycnophyidae) from 

San Juan Island, Washington, U.S.A. 

Andrey V. Adrianov and Robert P. Higgins 

(AVA) Institute of Marine Biology, Vladivostok 690041, Russia; 
(RPH) 2 Pond Lane, Asheville, North Carolina 28804, U.SA. 

Abstract. — A new species of kinorhynch, Pycnophyes parasanjuanensis, is 
described and illustrated from muddy sediments from a depth of 20 m at Friday 
Harbor, Washington (48°33'N, 123°04'W). It constitutes the second species of 
Pycnophyes from the San Juan Archipelago and the Pacific Coast of North 
America. The new species closely resembles P. sanjuanensis Higgins, 1961, 
the only other representative of this genus from this region. Like P. san- 
juanensis, P. parasanjuanensis has a wide, sculptured anterior margin of the 
first tergal plate and large circular zones of thin cuticle on the midsternal and 
episternal plates. It differs from the sympatric P. sanjuanensis and all other 
congeners in having longitudinal cuticular ridges near the lateral margins of 
the sternal plates and a fimbriate terminal border on segment 13. In addition, 
P. sanjuanensis differs from P. parasanjuanensis by its thick, robust lateral 
terminal spines with rounded tips. 



There have been few papers on Kino- 
rhyncha from the West Coast of the United 
States. Of the six species described, all but 
one have been from the San Juan Archi- 
pelago, located in the northwest of the state 
of Washington between Vancouver Island, 
Canada and the United States mainland. 
The first kinorhynch reported (Higgins 
1960) from this region was Echinoderes 
pennaki Higgins, 1960, a cyclorhagid, 
found at East Sound Bay of Orcas Island. 
The second publication (Higgins 1961) on 
kinorhyncha from this region described 
three species of homalorhagids, Pycnophyes 
sanjuanensis Higgins, 1961, Kinorhynchus 
ilyocryptus Higgins, 1961, and K. cata- 
phractus Higgins, 1961. Boykin (1965), in 
his unpublished dissertation, addressed the 
morphology of K. ilyocryptus. A second cy- 
clorhagid, Echinoderes kozloffi Higgins, 
1977 was the subject of a paper by Kozloff 
(1972) wherein he described the oviposition 
and hatching of the juvenile stage of this 
species described a few years later by Hig- 



gins (1977). This latter species was found 
in the intertidal zone of North Bay, San 
Juan Island. The only other kinorhynch de- 
scribed from the Pacific Coast of the United 
States is Echinoderes nybakkeni Higgins, 
1986. This, too, was described from the in- 
tertidal zone, from coarse beach sand at 
Carmel, California. 

Methods 

The five specimens upon which this 
study is based were collected by the senior 
author (AVA) on 21 Jul 1994. They were 
found in samples of mud taken by a 0.06 
m^ grab at a depth of 20 m, Friday Harbor, 
San Juan Island, San Juan Archipelago, lo- 
cated in the northwest section of the State 
of Washington between Vancouver Island 
and the United States mainland. Living ki- 
norhynchs were extracted from the sedi- 
ment by the "bubble-and-blot" method 
(Higgins 1983). Most specimens were fixed 
in 10% formalin. Some of were transferred 



VOLUME 109, NUMBER 2 



237 



to a glycerin-alcohol solution which was al- 
lowed to evaporate to glycerin. The glyc- 
erin-impregnated specimens then were 
mounted individually in Hoyer's-125 
mounting medium on slides for further ex- 
amination using phase-contrast and differ- 
ential interference contrast optics. A few 
specimens were selected for scanning elec- 
tron microscopic (SEM) study. These were 
transferred to a small tube, sealed with 42- 
|xm mesh nylon net, and placed in a small 
vessel of distilled water. Ethanol was added 
slowly until the contration was 100 percent; 
thereafter, the absolute ethanol was replaced 
several times. The tube and its contents 
were dried in a critical-point depression ap- 
paratus using carbon dioxide. Specimens 
were removed, mounted on SEM stubs and 
coated with gold-platinum. A Stereoscan 
Microscope 250 MK2 was used to study the 
specimens. 

Examination procedures followed the 
protocol described by Higgins (1983:4-7). 
Measurements are given in micrometers 
(|jLm). Ratios are expressed in percent of the 
total length (TL) measured on the midline, 
from the anterior margin of segment 3 (first 
trunk segment) to the posterior margin of 
segment 13, exclusive of spines. Maximum 
sternal width (MSW) is measured at the an- 
teroventral margin of the widest pair of 
sternal plates as first encountered in mea- 
suring each segment from anterior to pos- 
terior. Standard width (SE), or sternal width 
as segment 12, is measured at the antero- 
ventral margin of 12th sternal plates. The 
locality data from material examined are re- 
ferred to by the senior author's number 
(AVA). 

Two specimens of Pycnophyes parasan- 
juanensis, the holotypic female and allotyp- 
ic male have been deposited in the mei- 
ofaunal collection of the Institute of Marine 
Biology, Vladivostok, Russia. One paratype 
of P. parasanjuanensis is deposited in the 
Invertebrate Zoology collection of the Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C., 



U.S.A. under the catalog number USNM 

274223. 



Pycnophyes parasanjuanensis, new species 
Figs. 1-24 

Diagnosis. — Trunk length 740-800 jxm; 
trunk segments slightly increasing in width 
to segments 7-9, then tapering slightly at 
segments 11-12; subdorsal placids twice as 
wide as middorsal placids; anterior margin 
of first tergite slightly denticulate, shingled 
or reticulate, with long horn-like lateral pro- 
cesses 22 |xm long; posterior margin of ter- 
minal tergite clearly fimbriate; lateral ter- 
minal spines (LTS) 150-176 |xm long, 20- 
22% of trunk length; middorsal processes 
obtuse, on segments 4-11, nearly uniform 
in size, only slightly protruding beyond 
margin of tergite, each bearing 1-2 sensory 
setae; midsternal plate trapezoidal, anterior 
margin about 40% of posterior margin, an- 
terior border of midsternal plate projecting 
beyond anterior margins of episternites; 
posterior margins of segments 3-12 with 
longitudinal rows of minute spherical bod- 
ies, becoming less distinct posteriorly; pa- 
chycycli of segments 4-10(1 1) with unclear 
peg-and-socket articulation ventrally; 
anteromesial thickenings of ventral pachy- 
cycli prominent on segments 10-12 in fe- 
male and 9-12 in male, not adjacent at ven- 
tral midline; sternal plates of segments 4- 
1 2 with prominent cuticular ridges near lat- 
eral margins. 

Description. — Holotypic female (Figs. 1, 
2, 7-12), senior author's number AVA FH- 
1.18, (Figs. 1, 2, 7-12); Allotypic male, 
(Figs. 3-6, 15-20), senior author's number 
AVA FH-17; Note: data for allotypic male, 
if different from those of holotypic female, 
are in parenthesis and following those fe- 
male. TL 779 |xm (799 ixm), MSW-7 178 
ixm, 23% (22%) of TL; SW 156 |JLm (152 
|jLm), 20% (19%) of TL; LTS 156 fxm (176 
ixm), LTS/TL 20% (22%); middorsal pro- 
cesses on segments 4-11. 

Segment 1 : Head withdrawn in holotype. 



238 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 1, 2. Pycnophyes parasanjuanensis, holotypic female. 1, Ventral view; 2, Dorsal view. 



See mouth cone and nine oral syles (OS) in Segment 3: First trunk segment (Figs. 7, 

SEM photo of paratypic female. Fig. 21. 10), length 99 |xm (106 ixm); with lateral 

Segment 2: Not evident in holotype be- horn-like processes, 22 |jim long; anterior 

cause of withdrawn head. See neck placids margin finely denticulate, shingled or retic- 

(NP) in paratypic female. Fig. 22. ulate in appearance; pair of subdorsal tri- 



VOLUME 109, NUMBER 2 



239 




Figs. 3—6. Pycnophyes parasanjuanensis, allotypic male. 3, Segments 3, 4, dorsal view; 4, Segments 11- 
13, dorsal view; 5, Segments 3, 4, ventral view; 6, Segments 11-13, ventral view. 



angular muscle scars anterior to midlength 
of tergite and anterior to large scars of dor- 
soventral muscles; midsternal plate trape- 
zoidal (Figs. 5, 7, 15; note same in SEM 
photo of paratypic female, fig. 23), 88 |xm 
basal width tapering evenly to 33 |xm about 
one-sixth the distance from anterior margin 
and then becoming even, with round area 
of thin cuticle submarginally; anterior mar- 
gin of midsternal plate projecting beyond 



anteromesial margins of episternal plates; 
each episternite (Figs. 1, 5, 7, 15) with two 
adjacent areas of thinner cuticle, episternal 
plates with sensory seta near posterolateral 
margin; two lateral setae present. 

Segment 4: Length 78 fxm (80 |xm); ter- 
gite with two dorsolateral setae, middorsal 
process obtuse, with seta (male with seta on 
each side of middorsal process) pachycycli 
with peg-and-socket articulation ventrally; 



240 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 7-12. Pycnophyes parasanjuanensis, holotypic female. 7, Segments 3, 4, ventral view; 8, Segments 7, 
8, ventral view; 9, Segments 12, 13, ventral view; 10, Segments 3, 4, dorsal view; 11, Segments 7, 8, dorsal 
view; 12, Segment 12, dorsal view. All figures to same scale as Fig. 7. Abbreviations: CR, cuticular ridges; ES, 
episternal plate; LTS, lateral terminal spine; MP, middorsal process; MS, midsternal plate; RS, reticulate sculp- 
turing; SP, spermatophore remainder; s, prefix followed by segment number. 



VOLUME 109, NUMBER 2 



241 




Figs. 13, 14. Pycnophyes parasanjuanensis, paratypic female. 13, Segments 11—13, ventral view; 14, Seg- 
ments 11, 12, midventral thickenings. All figures to same scale as Fig. 7. Abbreviations: F, fimbriate margin of 
segment 13; MW, Midventral thickenings; SP, spermatophore remainder; SS, sensory seta; s, prefix followed by 
segment number. 



sternites with prominent cuticular ridges 
laterally and sub ventral setae; with two lat- 
eral setae. (Male with adhesive tube, 44 |xm 
long, anteromesial on each sternite. Figs. 5, 
16). 

Segment 5: Length 83 ixm (81 |xm); sim- 
ilar to segment 4 except for lack of lateral 
setae. 

Segment 6: Length 84 |xm; similar to 
segment 5 except for presence of two lateral 
setae. 

Segment 7: Length 89 |xm (87 ixm) (Figs. 
8, 17); similar to segment 6. 

Segment 8: Length 92 iJim (90 |xm); sim- 
ilar to segment 6 except for presence of two 
setae on middorsal process. 

Segment 9: Length 92 |xm (95 ixm); sim- 
ilar to segment 8 except for more prominent 
ventromesial pachycycli and only one seta 
on middorsal process. (Male with promi- 
nent anteromesial thickenings of ventral 
pachycycli.) 

Segment 10: Length 95 |xm (99 |xm); 
similar to segment 9 except for presence of 



well-developed anteromesial thickenings of 
ventromesial pachycycli (Figs. 1, 6, 14). 

Segment 11: Length 99 |xm (100 fxm); 
anteromesial thickenings longer and broad- 
er than in previous segment, similar to seg- 
ment 10 except for absence of lateral setae. 

Segment 12: Length 101 |jLm; no mid- 
dorsal process (Fig. 20); no dorsolateral se- 
tae; anteromesial thickenings longer and 
broader than in previous segment; sternites 
with ventrolateral seta; with lateral seta 
(Fig. 9; also note same segment shown in 
SEM photo of paratypic female, Fig. 24). 

Segment 13: Length 44 iJtm (43 jim); 
with two lateroterminal setae (Figs. 1, 4, 9, 
13); terminal border of tergite fimbriate 
Figs. 1, 2, 13); lateral terminal spines 156 
IJtm long. (Pair of penile spines, Figs. 6, 18, 
near anterolateral margin of each sternite, 
PS-1 48 iJtm long, PS-2 35 fxm long). 
(Small protuberance near lateroterminal 
margin of sternites). 

Sexual dimorphism. — The male differs 
from females in having prominent antero- 



242 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 15-20. Pycnophyes parasanjuanensis, allotypic male. 15, Segment 3, ventral view; 16, Segment 4, 
ventral view; 17, Segments 7-8, ventral view; 18, Segment 12-13, ventral view; 19, Segments 6, 7, dorsal view; 
20, Segments 11, 12, dorsal view. All figures to same scale as Fig. 7. Abbreviations: AT, adhesive tubes; ES, 
episternal plate; MP, middorsal process; MS, Midsternal plate; PS, penile spines; SS, sensory seta; s, prefix 
followed by segment number; TA, lateral terminal protubrances of segment 13; TS, thin cuticle area. 



VOLUME 109, NUMBER 2 



243 




Figs. 21-24. Pycnophyes parasanjuanensis, adult female, SEM photographs, scale equals 100 |xm. 21, Pro- 
truded mouth cone with oral styles, scale equals 10 p,m; 22, Segments 2-5, dorsal view, scale equals 100 |xm; 
23, Segments 2-5, ventral view, scale equals 100 |xm; 24, Segments 10-13, ventral view, scale equals 100 jjim. 
Abbreviations: CR, cuticular ridges; MP, middorsal process; MS, midsternal plate; NP, neck placid; OS, oral 
styles; RS, reticulate sculpturing; SS, sensory seta; S, sensory spot; SU, ectocommensal suctorian; s, prefix 
followed by segment number. 



244 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



mesial thickenings of ventral pachycycli on 
segments 9-12, two ventral adhesive tubes 
on segment 4, two lateroterminal protuber- 
ances on the margin of sternites of segment 
13, two pairs of penile spines (Figs. 6, 18) 
at the anterolateral margins of segment 13, 
and two sensory setae on the middorsal pro- 
cess of segment 4. 

Paratypic variation. — A single paratypic 
male, senior author's number AVA FH-1.16 
(Figs. 13, 14), TL 740 ixm; MSW-7 180 jxm 
(24% of TL), SW 156 jxm (21% of TL), 
LTS 152 |jLm (20.5% of TL). Paratypic fe- 
male differs from holotypic female in hav- 
ing two sensory setae on each middorsal 
process (Fig. 22, MP). 

Type material. — Holotype: adult female 
(AVA FH-1.18), allotype: adult male (AVA 
FH-1.17), paratype: adult female (AVA FH- 
1.16 (USNM 274223); Type Locality: Har- 
bor area south of Friday Harbor Laborato- 
ries, Friday Harbor, Washington, U.S.A. 
(48°33'N, 123°04'W); from mud at depth 
20 m, collected by A. V. Adrianov, 21 Jul 
94. 

Other material: Two specimens (Figs. 
21-24) mounted for SEM study, from type 
locality. 

Remarks. — Pycnophyes parasanjuanen- 
sis, n. sp. resembles only a few other con- 
geners. As it's name implies, it is similar to 
P. sanjuanensis Higgins, 1961. Both spe- 
cies have a wide reticulate-sculptured an- 
terior margin of the first tergite (Figs. 2, 3, 
1) (segment 3) and large round zones of 
thinner cuticle on the midsternal and epi- 
sternal plates of this same segment (Figs. 1, 
5, 7, 15). In addition, both are similar in 
size, general shape, shape of the midsternal 
plate and in the arrangement of setae and 
dorsal processes. However, P. parasanju- 
anensis is easily distinguished from the for- 
mer species by the shape of the lateral ter- 
minal spines (Figs. 1, 2, 4), by the presence 
of lateral cuticular ridges on the sternal 
plates (Figs. 1, 6, 8, 17, 24), and by the 
arrangement of anteromesial thickenings of 
ventral pachycycli. 

Other differences include the width of 



dorsal placids, and the shape of segment 13. 
The male of the new species bears latero- 
terminal protuberances on the caudal mar- 
gin of the sternites of the terminal segment 
(Figs. 6, 18). These are unique to this ge- 
nus, known otherwise in Kinorhynchus par- 
aneapolitanus (see Higgins & Adrianov 
1991). Two other members of this genus, P. 
dentatus Reinhard, 1881, and P. robustus 
Zelinka, 1928 from European waters, also 
have a wide sculptured anterior margin of 
the first tergite, but are distinguished by the 
arrangement and shape of middorsal pro- 
cesses and anteromesial thickenings of the 
ventral pachycycli. In contrast to the new 
species, P. dentatus has midventral thick- 
enings on segments 1 1 and 12 only, and has 
short lateral terminal spines. Pycnophyes 
robustus has midventral thickenings on seg- 
ments 8-12 and is further characterized by 
long lateral terminal spines, 34% of the 
trunk length, in contrast to P. parasanju- 
anensis (20% of the trunk length). The only 
other species of Pycnophyes having prom- 
inent cuticular ridges laterally on sternal 
plates is P. corrugatus Higgins, 1983. This 
species is distinguished from the new spe- 
cies by the absence of middorsal processes 
and shape of the areas of thin cuticle on the 
midsternal and epistemal plates. 

Pycnophyes parasanjuanensis is the sixth 
species of Kinorhyncha described from the 
northwest coast of the United States and 
from the San Juan Archipelago. It consti- 
tutes only the second member of the genus 
Pycnophyes described from the Northeast 
Pacific Ocean. 

Key to Adults of Pycnophyes 

1. Posterior margin of first tergite (seg- 
ment 3) with well-developed or subcu- 
ticular minute middorsal process 2 

— Posterior margin of first tergite always 
even, without middorsal process 18 

2. Middorsal process of first tergite round- 
ed or obtuse 3 

- Middorsal process of first tergite spi- 
nose, pointed 9 

3. Anteromesial thickenings of ventral pa- 



VOLUME 109, NUMBER 2 



245 



chycycli on segments 11-12, adjacent 

at ventral midline 

P. greenlandicus Higgins & Kristensen, 1988 

- Anteromesial thickenings of ventral pa- 
chycycli midventral thickenings prom- 
inent on other segments, adjacent or not 
adjacent at ventral midline 4 

4. Anteromesial thickenings of ventral 
pachycycli on segments 6-12 5 

- Anteromesial thickenings of ventral pa- 
chycycli on segments 9-12 6 

5. Anteromesial thickenings of ventral pa- 
chycycli narrowly elongate, most of 
them longer than half the sternite 
length, lateral terminal spines about 
25% of trunk length 

. . . P.canadensisVixggms&Koxczyns\d,\9%9 

- Anteromesial thickenings of ventral pa- 
chycycli shorter than half of sternite 
length, lateral terminal spines about 

10-15% of trunk length 

P. communis Zelinka, 1928 

6. Posterior margin of sternite 13 with two 
midventral conical protrusions extend- 
ing to the margin of the tergite 

P. mokievskii Adrianov 1995 

- Posterior margin of sternite 13 without 
midventral conical protrusions 7 

7. Posteromesial ventral pachycycli of 
segment 12 prominent, deeply incised 
anteriorly or elongated posteriorly . . 

P. spitsbergenensis Adrianov 1995 

- Posteromesial ventral pachycycli not 
prominent, underdeveloped, not in- 
cised, not elongated posteriorly 8 

8. Middorsal processes minute, on seg- 
ments 3-8(9) . . P. maximus Reimer, 1963 

- Middorsal processes broadly rounded, 
on segments 3-12 

. . P. borealis Higgins & Korczynski 1989 

9. Patches of puncations near lateral mar- 
gins of sternal plates on segment 4-12 

P. iniorhaptus Higgins, 1983 

- Patches of punctations near lateral mar- 
gins of sternal plates on segments 4-12 
absent 10 

10. Middorsal spinous process on segment 
12 long, extending well beyond terminal 
margin .... P. chukchiensis Higgins, 1991 

- Middorsal spinous process on segment 

1 2 absent or poorly developed 11 

1 1 . Anterior margin of first tergite with wide 
area of cuticular netting or mosaic pat- 



tern; sternal plates of segment 12 with 
vertical cuticular striations near lateral 
margins P. dentatus Zelinka, 1928 

- Anterior margin of first tergite without 
netting or mosaic pattern; sternal plates 
of segment 12 without cuticular stria- 
tions near lateral margins 12 

12. Thin area of cuticle at anteromesial 
margin of episternal plates double or 
longitudinally divided 13 

- Thin area of cuticle at anteromesial 
margin of episternal plates single or ab- 
sent 14 

13. Thin area of cuticle at anteromesial 
margin of episternal plates elongated, 
about 33% of plate length; anteromesial 
thickenings of ventral pachycycli of 
segment 12 widely separated; lateral 
terminal spines longer than width of 
segment 12 . . .P. flaveolatus Zelinka, 1928 

- Thin area of cuticle at anteromesial 
margin of episternal plates short, less 
than 20% of plate length; anteromesial 
thickenings of ventral pachycycli of 
segment 12 adjacent at ventral midline; 
lateral terminal spines shorter than 
width of segment 12 

P. calmani Southern, 1914 

14. Thin area of cuticle at anterior margin 
of midsternal plate strongly flattened or 
oval-shaped, round on episternal plates; 
middorsal processes minute, barely pro- 
truding beyond posterior margin of ter- 
gite; anteromesial thickenings of ven- 
tral pachycycli on segments 11-12 . . 

P. frequens Blake, 1930 

- Thin area of cuticle at anterior margin 
of midsternal and episternal plates un- 
derdeveloped; middorsal processes 
elongated; anteromesial thickenings of 
ventral pachycycli, if present, only on 
segment 12 15 

15. Segment 2 with 3 dorsal placids .... 

P. odhneri Lang, 1949 

- Segment 2 always with 4 dorsal placids 

16 

16. Anterior margin of midsternal plate 
projecting well beyond anteromesial 
margins of episternal plates; patches of 
punctations at middorsal processes of 
tergites P. carinatus Zelinka, 1928 

- Anterior margin of midsternal plate 
even with, not projecting beyond, an- 



246 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



teromesial margins of episternal plates; 
no patches of punctations at middorsal 
processes of tergites 17 

17. Posterior margin of midsternal plate 
twice the width of anterior margin . . 

P. chiliensis Lang, 1953 

- Posterior margin of midsternal plate 
only slightly wider than anterior margin 

P. cryopygus Higgins & ICristensen, 1988 

18. Anterior margin of first tergite scallo- 
ped, with one middorsal and two dor- 
solateral projections, anterior margin 
between projections concave 19 

- Anterior margin of first tergite dentate, 
coronate (at least seven or more projec- 
tions) or even 20 

19. Tergal plates of segments 4, 5 with 
middorsal processes 

P. ponticus Reinhard, 1881 

- Tergal plates of segments 4, 5 without 
middorsal processes 

P. kielensis Zelinka, 1928 

20. Anterior margin of first tergite coro- 
nate, with prominent middorsal projec- 
tion and three lateral projections on 
each side, margin between projections 
denticulate P. rugosus Zelinka, 1928 

- Anterior margin of first tergite even or 
evenly dentate without projections ... 21 

21. Anterior margin of midsternal plate 
very narrow, about 25% of posterior 
margin P. ecphantor Higgins, 1983 

- Anterior margin of midsternal plate rel- 
atively broad, about 33-50% of poste- 
rior margin 22 

22. Anterior margin of first tergite with 
wide area of reticulate, net-or-mosaic- 
like pattern 23 

- Anterior margin of first tergite without 
wide area of reticulate, net-or-mosaic- 
like pattern 24 

23. Lateral terminal spines thick, robust 
and obtuse (not pointed at top); anter- 
omesial thickenings of ventral pachy- 
cycli on segments 8-12 

P. sanjuanensis Higgins, 1961 

- Lateral terminal spines pointed at top; 
prominent anteromesial thickenings of 
ventral pachycycli on segments 10-12 
only ... P. parasanjuanensis, new species 

24. Thin area of cuticle at anterior margin 
of midsternal plate double, divided into 



two separated areas 

P. tubuliferus Adrianov, 1989 

- Thin area of cuticle at anterior margin 

of midsternal plate single or absent . . 25 

25. Lateral terminal spines nearly equal to 
length of segment 12 

P. zelinkaei Southern, 1914 

- Lateral terminal spines longer than 
combined length of segments 12 and 13 

26 

26. Thin area of cuticle at anteromesial 
margin of episternal plates double, di- 
vided into two large separated areas: 
anteromesial thickenings of ventral pa- 
chycycli not prominent on any segment 

P. sculptus Lang, 1949 

- Thin area of cuticle at anteromesial 
margin of episternal plates, if present, 
not double; anteromesial thickenings of 
ventral pachycycli present 27 

27. Anteromesial thickenings of ventral pa- 
chycycli thin, narrowly elongate, on 
segments 8-12; anterior margin of first 
tergite pectinate 

P. robustus Zelinka, 1928 

- Anteromesial thickenings of ventral pa- 
chycycli not narrowly elongate, on oth- 
er segments; anterior margin of first ter- 
gite even or slightly denticulate 28 

28. Thin area of cuticle at anterior margin 

of midsternal plate present 29 

- Thin area of cuticle at anterior margin 

of midsternal plate absent 31 

29. Thin area of cuticle at anteromesial 
margin of episternal plates present; pos- 
terior margin of terminal tergite without 
lateral bulbous protrusions 30 

- Thin area of cuticle at anteromesial 
margin of episternal plates absent; pos- 
terior margin of terminal tergite with 
lateral bulbous protrusions 

P. emarginatus Higgins, 1983 

30. Sternal plates of segments 11, 12 with 
strong longitudinal cuticular ridges near 
lateral margins; anterior margins of ter- 
gal and sternal plates without wide ar- 
eas of cuticular microrelief 

P. corrugatus Higgins, 1983 

- Sternal plates of segments 11, 12 with- 
out cuticular ridges; anterior margins of 
tergal and sternal plates with wide areas 
of cuticular microrelief 

P. egyptensis Higgins, 1966 



VOLUME 109, NUMBER 2 



247 



31. Thin area of cuticle at anteromesial 
margin of episternal plates present; an- 
teromesial thickenings of ventral pa- 
chycycli on segments 10-12 

P. longicornis Higgins, 1983 

- Thin area of cuticle at anteromesial 
margin of episternal plates absent; an- 
teromesial thickenings of ventral pa- 
chycycli on segment 12 

P. beaufortensis Higgins, 1964 



Acknowledgments 

The senior author wishes to thank Dr. D. 
A. Willows, Director, Friday Harbor Labo- 
ratories, University of Washington for the 
opportunity of working at this facility. Ac- 
knowledgement is also made to the Grass 
Foundation and their generous award of a 
Post-doctoral Fellowship which made this 
study possible. We are grateful to the Sum- 
ner Gerard Foundation for providing pub- 
lication funds. 

Literature Cited 

Adrianov, A. V. 1989. The first report of Kinorhyncha 
of the Sea of Japan. — Zoologichesky Zhurnal 
61:17-27 (in Russian with English summary). 

. 1995. The first description of kinorhynchs 

from the Spitsbergen Archipelago (Greenland 
Sea) with a key to the genus Pycnophyes (Hom- 
alorhagida, Kinorhyncha). — Canadian Journal 
of Zoology 73:1554-1566. 

Blake, C. H. 1930. New species of worms belonging 
to the order Echinodera.— Biological Survey of 
the Mount Desert Region 4:3-8. 

Boykin, J. C. 1965. The anatomy of Trachydemus il- 
yocryptus (Kinorhyncha). Unpublished M.S. 
thesis, University of Washington, Seattle, 94 pp. 

Higgins, R. R 1960. A new species of Echinoderes 
(Kinorhyncha from Puget Sound. — Transactions 
of the American Microscopical Society 79:85- 
91. 

— . 1961. Three new homalorhagid kinorhynchs 

from the San Juan Archipelago, Washington. — 
Journal of the Elisha Mitchell Scientific Society 
77:81-88. 

. 1964. Three new kinorhynchs from the North 

Carolina Coast. — Bulletin of Marine Science of 
the Gulf and Caribbean 14:479-493. 

. 1966. Faunistic studies in the Red Sea (in 



winter, 1961-1962). Part II. Kinorhynchs from 
the area of Al-Ghardaqa. — Zoologisches Jahr- 
biicher. Systematic Oekologie und Geographic 
der Tiere 93:118-126. 

— . 1977. Redescription of Echinoderes dujar- 
dini (Kinorhyncha) with descriptions of closely 
related species. — Smithsonian Contributions to 
Zoology 248:1-26. 

— . 1983. The Atlantic Barrier Reef ecosystem 
at Carrie Bow Cay, Belize. II. Kinorhyncha. — 
Smithsonian Contributions to Marine Science 
18:1-131. 

— . 1986. A new species of Echinoderes (Kino- 
rhyncha: Cyclorhagida) from a coarse-sand Cal- 
ifornia Beach. — Transactions of the American 
Microscopical Society 105:66-73. 

— . 1991. Pycnophyes chukchiensis, a new hom- 
alorhagid kinorhynch from the Arctic Sea. — 
Proceedings of the Biological Society of Wash- 
ington 104:184-188. 

— , & A. V. Adrianov. 1991. Kinorhyncha from 
the Black Sea. I. Redescription of Kinorhynchus 
paraneapolitanus. — Transactions of the Ameri- 
can Microscopical Society 110:328-336. 

— , & R. M. Kristensen. 1988. Kinorhyncha from 
Disko Island, West Greenland. — Smithsonian 
Contributions to Zoology 438:1-70. 

— , & R. E. Korczynski. 1989. Two new species 



of Pycnophyes (Homalorhagida, Kinorhyncha) 
from the Canadian coast of the Beaufort Sea. — 
Canadian Journal of Zoology 67:2056-2064. 

Kozloff, J. N. 1972. Some aspects of development in 
Echinoderes Kinorhyncha). — Transactions of 
the American Microscopical Society 91:119- 
130. 

Lang, K. 1949. Echinoderida. Pp. 1-8 in N. H. 
Odhner, ed.. Further zoological results of the 
Swedish Antarctic Expedition, 1901-1903, Vol- 
ume 4, Number 2. C. W. K. Gleerup, Lund, 22 
pp. 

. 1953. Reports of the Lund Unniversity Chile 

Expedition 1948-49. 9. Echinoderida.— Kun- 
glia Fysiografiska Sallskapets Handlingar, N. F 
64:1-8. 

Reimer, L. 1963. Zur Verbreitung der Kinorhyncha in 
der mittleren Ostsee. — Zoologischer Anzeiger 
171:440-447. 

Reinhard, W. 1881. Uber Echinoderes und Desmos- 
colex der Umgegend von Odessa. — Zoologisch- 
er Anzeiger 4:588-592. 

Southern, R. 1914. Nemathelmia, Kinorhyncha and 
Chaetognatha. in Clare Island Survey, Part 
54. — Proceedings of the Royal Irish Academy 
31:1-80. 

Zelinka, C. 1928. Monographic der Echinodera. Wil- 
helm Engelmann, Leipzig, 396 pp. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 

109(2):248-255. 1996. 

A new genus and species of ampharetid polychaete from deep-sea 
hydrothermal vent community in the Azores triple-junction area 

Daniel Desbruyeres and Lucien Laubier 

(DD) Departement Environnement Profond, Centre de Brest de I'lFREMER, 

B.P. 70, 29280 Plouzane, France; 

(LL) Laboratoire de Physiologic dcs Etrcs marins, Institut Oceanographique, 

195 rue Saint Jacques, 75005 Paris, France 

Abstract. — Amathys lutzi, a new genus and species of Ampharetinae (Poly- 
chaeta: Ampharetidae), is described from active hydrothermal vents on the 
Lucky Strike segment, close to the Azores triple-junction area of the Mid- 
Atlantic Ridge. It is found in the same micro-habitat than Amphisamytha gal- 
apagensis Zottoli, 1983. It differs from all previously known Ampharetinae by 
the number (17) of thoracic uncinigerous segments. 



In 1993, the Lucky Strike expedition 
(cruise Atlantis II 129-6) visited an active 
hydrothermal field situated at 37°17.538'N 
and 32° 1 6.490' W in 1626 m. The expedi- 
tion consisted of six dives of the DSRV Al- 
vin and aimed to find and describe active 
hydrothermal sites and the biological com- 
munities associated with venting areas. 
Based on this first exploration, the Diva 2 
expedition (1994) completed 24 dives of 
the submersible Nautile on the same hydro- 
thermal area. Hydrothermal activity occurs 
around a lava lake situated in the depression 
between three cones which form the sum- 
mit of a seamount, at the center of a long 
broad ridge segment that extends from 
37°00'N to 37°35'N; hydrothermal activity 
includes flanges with pool temperature 
<200°C to black smokers venting fluids as 
hot as 333°C (Langmuir et al., 1993, pers. 
obs.). The dominant organisms in these hy- 
drothermal fields are populations of a new 
species of mussel (bathymodiolid) that col- 
onizes hydrothermal sulfide edifices and 
their vicinity. Other less conspicuous com- 
ponents of the fauna include new species of 
alvinocarid shrimps, peracaridan crusta- 
ceans, bythograeid crabs, sea-urchins and 
several species of polychaetes including a 
polynoid commensal of mussels (Van Do- 



ver et al. 1993). Ampharetid polychaetes 
were sorted from mussel washings, or 
found in tubes attached to mussel shell 
hinges and to sulfide or basaltic rocks. 

Materials and Methods 

The specimens were collected using Al- 
vin and Nautile manipulators. The samples 
were brought to the surface inside insulated 
boxes and were washed and sieved through 
a 250-|JLm mesh. They were sorted partially 
aboard the mother-vessels, then fixed in 
buffered formalin and preserved in 80% 
ethanol after a one day fixation. In the lab- 
oratory, specimens are dehydrated to ethyl 
alcohol absolute, critical point dried, put on 
stubs and coated with gold. Finally, the 
specimens were examined using a Philips 
XL 30 scanning electron microscope. 

Order Terebellida 

Family Ampharetidae Malmgren, 1865 

Subfamily Ampharetinae Chamberlin, 

1919 

Amathys, new genus 

Diagnosis. — Thorax with 20 setigerous 
segments, of which the last 17 are uncini- 
gerous. Abdomen tapering rapidly with 16- 
20 uncinigerous segments, no notopodial or 



VOLUME 109, NUMBER 2 



249 



neuropodial cirri or vestigial abdominal no- 
topodia. Notopodia of the first setigerous 
segment reduced. Four pairs of smooth 
branchiae. A buccal ciliated membrane 
bearing dorsally clavate and grooved ten- 
tacles. Anus terminal without cirri. No 
modified segment. 

Genus. — Amathys 

Type species. — Amathys lutzi 

Etymology. — Amathys is an anagram of 
Samytha. Gender is male. 

Remarks. — This genus differs from all 
known genera of Ampharetinae by having 
17 uncinigerous thoracic segments. 

Amathys lutzi, new species 
Figs. 1, 2, 3 

Etymology. — This species is dedicated to 
Dr. Richard A. Lutz from Rutgers Univer- 
sity, New Jersey, USA in an attempt to ex- 
press our friendly thanks for several oppor- 
tunities that he gave us to dive with the 
DSRV Alvin in several Eastern Pacific hy- 
dro thermal vent areas. 

Material. — Lucky Strike expedition: dive 
Alvin 2606, 1 June 1993 (M. Tivey, L. Sal- 
danha, observers): 5 specimens, dive Alvin 
2607, 2 June 1993 (C. Van Dover, T. Em- 
erson, observers): 1 specimen. Diva 2 ex- 
pedition: dive Nautile 912/40-01, 3 June 
1994 (A.M. Alayse, observer): 1 specimen; 
dive Nautile 913/41-02, 4 June 1994 (R 
Crassous, observer): 15 specimens; dive 
Nautile 915/43-04, 6 June 1994 (D. Des- 
bruyeres, observer): 2 specimens (including 
holotype); dive Nautile 916/44-05, 7 June 
1994 (E Barriga, observer): 117 specimens; 
dive Nautile 917/45-06, 8 June 1994 (L. 
Saldanha, observer): 45 specimens. All 
specimens from different vent sites within 
the Lucky Strike area (latitude from 
37° 17,25' to 37° 1 7,63 'N, longitude from 
32°16,50' to 32°16,90'W, depth from 1622 
to 1725 m). All specimens from washings 
of mussel clumps or tubes attached to mus- 
sel shells and sulfide or basaltic rocks. Ho- 
lotype (USNM 170025), Paratypes 
(MNHN, Paris, UD 851/A928). 



Description. — Body whitish, without 
color pattern when preserved. Holotype 
complete, about 14.8 mm in length and 4.3 
mm in greatest width, with 38 setigerous 
segments. Size of paratypes from 1 .25 mm 
to 15.7 mm in length and from 0.25 mm to 
4.4 mm in width. Paratype used for SEM 
observations complete, 6 mm long and 2.1 
mm wide. Prostomium indistinctly trilobed, 
lacking glandular ridges and eye spots but 
bearing two transversal nuchal slits under- 
lined by secretions and mineral particles 
(Fig. 2a). Buccal tentacles smooth, entire, 
clavate, grooved and ciliated (Fig. 2b), in- 
serted on the latero-dorsal part of a folded 
"feeding" membrane heavily ciliated (Fig. 
2c) and sprinkled with organic and mineral 
aggregates. First two segments (I and II ac- 
cording to Day 1964) achaetous and fused, 
forming the lower lip. Segment III lacking 
palae. Thorax with twenty setigerous seg- 
ments, the last seventeen being unciniger- 
ous. The first setiger, segment IV, with a 
vestigial notopodium and reduced bundles 
of capillary setae. Four pairs of smooth, cy- 
lindrical branchiae on dorsal surface of seg- 
ments III-VI, all similar, regularly attenu- 
ated. Branchial length about % of the body 
length in preserved specimens. Branchiop- 
hores short and not fused. Small mid-dorsal 
gap between the two branchial groups but 
no web. The two outer branchiophores in- 
serted between segments III and IV, the two 
inner branchiophores in segments V and VI. 
Notopodia increasing in size from setiger- 
ous segments 2 to 5. Notopodium cylindri- 
co-conical, slightly flattened anterio-poste- 
riorly with two series of winged capillary 
setae (one short, the other approximately 
250 [xm long). Notosetae capillary slightly 
unilimbated, covered with continuous layer 
of minute spinelets when viewed under 
scanning electron microscope (Fig. 3a, b). 
Neuropodial lobes (uncinigerous pinnules) 
from setiger 4, each with a single row of 
toothed uncini, tips oriented forward (23- 
30 uncini per row). 17 uncinigerous tori on 
thoracic segments and 16-20 uncinigerous 
pinnules on abdominal segments. Thoracic 



250 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 1. Amathys lutzi, new genus, new species. Entire animal (holotype) in ventro-lateral view. 



VOLUME 109, NUMBER 2 



251 




Fig. 2. Amathys lutzi, new genus, new species, a, prostomium in dorso-lateral view (left side), b, ciliated 
groove of a buccal tentacle, c, detail of the ciliated surface of the fold, buccal membrane sprinkled with organic 
and mineral aggregates, d, thoracic uncinus in lateral view, e, abdominal parapodia in ventro lateral view (left 
side of the body), f, thoracic uncini with a duplication of the upper tooth, g, tube. 



252 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




o 

a 
o 






VOLUME 109, NUMBER 2 



253 



parapodia without cirri. Each thoracic un- 
cinus with a single row of four teeth (Fig. 
2d), the upper ones, sometimes but seldom, 
laterally duplicated (Fig. 2f). Abdominal 
uncini with same shape and size as thoracic 
ones. Small teeth present at the base of the 
main ones laterally on thoracic and abdom- 
inal uncini. A rounded glandular pad above 
each abdominal uncinigerous pinnule (Fig. 
2e). Abdominal notopodial lobes absent. 
Abdomen short (one-third of the body 
length). Pygidium terminal without circle of 
papillae or anal cirri. No indication of sex- 
ually mature specimens. 

Tube. — Mucus lined tubes covered with 
rusty colored mineral particles, agglomer- 
ated with mussel periostracum or byssal 
threat fragments as well as pieces of gas- 
tropod shells; about three times worm 
length (Fig. 2g). Sometimes, tube simple 
covered with gray mud. 

Discussion. — The morphology of this 
species is reminiscent of that of Amphisa- 
mytha galapagensis Zottoli, 1983 and that 
of Amphisamytha fauchaldi Solis-Weiss & 
Hernandez- Alcantara 1994, both found 
within the Pacific hydrothermal vent envi- 
ronment. In the Atlantic Ocean, McHugh & 
Tunnicliffe (1994) report, the presence of 
Amphisamytha galapagensis listed as "Am- 
pharetidae gen. sp." by Segonzac (Segon- 
zac 1992); we do not agree with this as- 
sumption and we here confirm, after dis- 
cussion with M. Segonzac, that all speci- 
mens (including Segonzac's specimens) 
from both Snake Pit (M.A.R., 23°20'N) and 
Broken Spur (M.A.R., 29°10'N) vent fields 
were damaged and macerated and that their 
bad shape doesn't allow determination, 
even at the generic level. 

Amathys lutzi, new species, clearly be- 
longs to a group of species having in com- 
mon a ciliated buccal membrane, grooved 
ciliated or entire clavate buccal tentacles, 
no palae, reduced setation on segment IV. 
This group gathers species having three or 
four pairs of branchiae, different numbers 
of thoracic uncinigerous segments, one or 
two rows of teeth on the thoracic unicini. 



This collective group gathers together Am- 
phisamytha galapagensis Zottoli, 1983, 
Amphisamytha fauchaldi Solis-Weiss & 
Hernandez- Alcantara, 1994, Amythas mem- 
branifera Benham, 1921 and Samytha cal- 
iforniensis Hartman, 1969. A. lutzi differs 
from them by the presence of 17 uncini- 
gerous thoracic segments instead of 14 in 
A. galapagensis and A. fauchaldi and the 
shape of abdominal uncini of the anal part 
in the former species (Zottoli 1983). This 
high number of uncinigerous thoracic seg- 
ments is unusual among Ampharetinae; un- 
til the present work it varied within the sub- 
family from a minimum of 9 {Mugga and 
Egamella) to a maximum of 15 (Weddel- 
lia). Since Day's (1964) revision, the num- 
ber of thoracic uncinigerous segments has 
been used by taxonomists as a character 
that is consistent at the generic level. 

Day's revision was justified by the high 
number of genera (27) and the difficulty of 
the recognition of genera. Day (1964) pro- 
posed to use the number of gills, the num- 
ber of thoracic uncinigerous segments, the 
presence of a glandular ridge on the prosto- 
mium and the presence of notopodial cirri 
as generic characters. Since then, 24 new 
genera were described, all of them still re- 
maining monospecific (see Holthe 1986 for 
review, Jirkov 1986, Solis-Weiss 1993). In 
1979, using multi variable analysis on a phe- 
netical matrix based on Day's taxonomic 
characters, Chardy & Desbruyeres (1979) 
demonstrated that these characters are only 
discriminant among a few major shallow 
water genera like Amphicteis, Ampharete, 
Amage or Anobothrus but that they are 
poorly discriminant in the case of most of 
the abyssal species which look as a contin- 
uum when using Day's characters. What- 
ever should be the present generic cluster- 
ing for Ampharetid species, it suffers deep- 
ly from the history of sampling which start- 
ed in shallow waters where evolutive 
radiation is limited to a few rather well de- 
fined genera. Sampling of the deep-sea ben- 
thos, which increased rapidly since the 
Challenger expedition, brought numerous 



254 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



new forms whose morphological patterns 
do not fall inside the shallow water model 
variations. In deep-sea species, the different 
generic characters as cited above evolve in- 
dependently, thus leading to a continuum of 
species where generic patterns are difficult 
to recognize either by aggregation or by di- 
chotomy (Chardy & Desbruyeres op. cit.). 
Unfortunately, most of the new taxa were 
described on the basis of unique specimens 
that were often truncated and sometimes 
damaged during recovery of samples and 
sieving of abyssal sediment, which led to a 
difficult situation in the scope of a new re- 
vision. 

In his recent work on Terebellomorpha, 
Holthe (1986) proposed to gather Samytha, 
Samythella, Eusamytha, Amythas, Demun- 
ciger and Alkmaria in a new tribe named 
Samythini for Ampharetinae with tentacles 
smooth, no glandular ridge in the prosto- 
mium. Most of these genera have a rather 
developed buccal membrane as does 
Amathys lutzi. Surprisingly, Holthe placed 
Amphisamytha among "Ampharetinae with 
uncertain tribal affinities" when the diag- 
nosis of this genus fits well with the char- 
acteristics of the tribus Samythini. I propose 
here to include Amphisamytha and Amathys 
inside Samithini. 

The erection of the new genus Amathys 
is a conservative position which takes into 
account the classical taxonomic characters 
as displayed in Day's revision (1964). In 
that context, the erection of a new genus is 
fully justified, but, as mentioned by Holthe 
(1986), ". . . when all genera become 
monotypic, the generic category has be- 
come void of information." Full phyloge- 
netic analyses of morphological and molec- 
ular characters may lead to major generic 
revisions of the Ampharetinae in future. 

Acknowledgments 

We thank Philippe Crassous (scanning 
electron microscope) and Violaine Martin 
(drawings) for technical assistance, Patrick 
Briand (CENTOB) for sorting specimens 



and Dr. C. Langmuir, chief scientist of the 
Lucky Strike expedition, who kindly invit- 
ed one of us (DD) to participate to the 
cruise and to dive with the DSRV Alvin. 

Literature Cited 

Benham, W. B. 192L Polychaeta. — Scientific Re- 
ports, Australian Antarctic Expedition 1911-14. 
Sen C, 6(3): 1-128. 

Chardy, P., & D. Desbruyeres. 1979. La classification 
multicritere. Application a la revision de la 
sous-famille des Ampharetinae (Annelides Pol- 
ychetes). — Annales de Biologic 18(11-12): 
521-537. 

Day, J. H. 1964. A review of the family Ampharetidae 
(Polychaeta). — Annals of the South African 
Museum 48(4):97-120. 

Hartman, O. 1969. Atlas of the sedentariate poly- 
chaetous annelids from California. — Allan Han- 
cock Foundation, Los Angeles, 812 pp. 

Holthe, T. 1986. Evolution, systematics, and distri- 
bution of the Polychaeta Terebellomorpha, with 
a catalogue of the taxa and a bibliography. — 
Gunneria 55:1-236. 

Jirkov, I. 1986. Zatsepina rittichae gen. sp. n. (Poly- 
chaeta, Ampharetidae) from the Norwegian and 
Barents seas. — Zoologichesky Zhurnal 65(3): 
325-332. 

Langmuir, C. H., et al. 1993. Geological setting and 
characteristics of the Lucky Strike vent field at 
37°17'N on the Mid-Atlantic Ridge. — Eos 
Transactions of the American Geophysical 
Union 74:99. 

McHugh, D., & V. Tunnicliffe. 1994. Ecology and 
reproductive biology of the hydrothermal vent 
polychaete Amphisamytha galapagensis (Am- 
pharetidae). — Marine Ecology Progress Series 
106:111-120. 

Segonzac, M. 1992. Les peuplements associes a 
I'hydrothermalisme oceanique du Snake Pit 
(dorsale medio-atlantique; 23°N, 3480 m): com- 
position et microdistribution de la mega- 
faune. — Comptes Rendus Academic Sciences 
Paris, Serie III 314:593-600. 

Soils- Weiss, V. 1993. Grass leia hydrothermalis, a 
new genus and species of Ampharetidae (An- 
nelida: Polychaeta) from the hydrothermal vents 
off the Oregon coast (U.S.A.), at Gorda 
Ridge. — Procedings of the Biological Society of 
Washington 106:661-665. 

, & P. Hernandez- Alcantara 1994. Amphisa- 
mytha fauchaldi: A new species of ampharetid 
(Annelida: Polychaeta) from the hydrothermal 
vents at Guaymas basin, Mexico. — Bulletin of 
Southern California Academy of Sciences 93: 
127-134. 

Van Dover, C. L., D. Desbruyeres, L. Saldanha, A. 



VOLUME 109, NUMBER 2 . 255 

Fiala-Medioni, & C. Langmuir. 1993. A new new species of ampharetid polychaete from the 

faunal province at the Lucky Strike hydrother- vicinity of abyssal hydrothermal vents in the 

mal vent field. — Eos Transactions of the Amer- Galapagos rift, and the role of this species in 

ican Geophysical Union 74:100. rift ecosystem. — Proceedings of the Biological 

Zottoli, R. A. 1983. Amphisamytha galapagensis a Society of Washington 96:79-391. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):256-263. 1996. 

Description of a new branchiobdellidan species, with observations on 

three other species, and a key to the genus Pterodrilus 

(Annelida: Clitellata) 

Stuart R. Gelder 

Department of Science, University of Maine at Presque Isle, 181 Main Street, 
Presque Isle, Maine 04769-2888, U.S.A. 

Abstract. — Pterodrilus annulatus, a new species, is described from Tennes- 
see, and compared with the west coast species Cambarincola serratus. An 
emended diagnosis of Pterodrilus and new observations on the anatomy of P. 
cedrus, P. choritonamus and P. hobbsi are presented. A key to the species of 
the genus, based on external segmental features, is given. 



Information on the branchiobdellidan ge- 
nus Pterodrilus was reviewed and extended 
by Holt (1968). The genus appears to be 
restricted to the eastern United States and 
its phylogenetic relationship with other gen- 
era was discussed in a synopsis of the Bran- 
chiobdellida (Holt 1986). More recently 
Pterodrilus was included in a checklist of 
North and Central American cambarincol- 
ids prepared by Holt & Opell (1993). 

Collections of the crayfish, Orconectes 
placidens (Hagen, 1870), from a single site 
on the Harpeth River south of Franklin, 
Tennessee, yielded eight species of bran- 
chiobdellidans. Usually, at least six of these 
species were present on a single host. Live 
branchiobdellidans were relaxed in Perrier 
Water, fixed in formalin-ethanol-acetic acid 
(FA A), dehydrated in a graded series of eth- 
anohwater solutions, cleared in methyl sa- 
licylate, and mounted on a microscope slide 
in Canada balsam (Brinkhurst & Gelder 
1991). The anatomical nomenclature used 
in this paper follows that described in 
Brinkhurst & Gelder (1991) with two ex- 
ceptions. The spermi ducal gland and ejac- 
ulatory duct are now referred to as the glan- 
dular atrium and muscular atrium respec- 
tively so that the terminology becomes con- 
sistent with that used in the rest of the 
Clitellata. Specimens were examined from 



the collections of Perry C. Holt (PCH), the 
National Museum of Natural History, 
Smithsonian Institution (USNM) and the 
author (SRG). 

During the process of identifying the 
branchiobdellidans from the Harpeth River, 
a new species of Pterodrilus was recog- 
nized. To confirm the new species status, 
type specimens of four species of Pterod- 
rilus and Cambarincola serratus Holt, 1981 
were borrowed from the National Museum 
and examined. These studies resulted in a 
clarification of the published descriptions of 
some of the anatomical features in certain 
pterodrilids, and the correction of some pre- 
viously identified specimens. The keys con- 
structed by Hoh (1968) and Holt & Opell 
(1993) for identifying species of Pterodri- 
lus utilize both external and internal ana- 
tomical features. However, following my 
examinations of the literature and speci- 
mens, a key was devised that used only ex- 
ternal anatomical features. This key should 
greatly simplify the identification of both 
living and preserved specimens for biolo- 
gists interested in the genus Pterodrilus. 

Pterodrilus Moore, 1895 

References. — Moore 1895:449-456, 
Goodnight 1940:58, Holt 1968:1, 3, 1986: 
698. 



VOLUME 109, NUMBER 2 



257 



Diagnosis. — Single anterior nephridial 
pore; vasa deferentia entering glandular 
atrium entally; body less than 2.0 mm long; 
dorsal ridge across segment 8, other seg- 
ments may have ridges and additional fin- 
ger-like projections; dental formula 5/4; 
prostate gland arises from ectal to mid re- 
gion of glandular atrium, and incompletely 
separated from it; penis muscular, protrusi- 
ble; spermatheca present (emended by 
SRG). 

Pterodrilus annulatus, new species 
Fig. lA-F, 3G 

Type specimens. — Holotype, USNM 
169996, and two paratypes, USNM 169997, 
and 169998 on Orconectes placidens (Ha- 
gen) removed from the Harpeth River, 6 
miles SE of Franklin, Williamson Co., Ten- 
nessee (35°59'N, 86°32'W) by S. R. Gelder 
and R. O. Brinkhurst, 17 Oct 1990. 

Diagnosis. — Body about 1.0 mm long, 
very prominent transverse ridges across the 
dorsal and lateral surfaces of segments 2 
through 8, the last is fan-like, supernumer- 
ary muscles present, no dorsal segmental 
appendages; dorsal lip four slight rises; oral 
papillae 16; jaws small, lightly sclerotized, 
dorsal kidney-shaped, ventral quadrangular, 
dental formula 5/4; glandular atrium sub- 
cylindrical, length about V2 and width Va di- 
ameter of segment; prostate gland length 
and diameter subequal to glandular atrium, 
differentiated, ental bulb absent; muscular 
atrium length about V2 glandular atrium; 
bursa ovoid, length about Vs segment di- 
ameter, penial sheath in ental % of bursa; 
spermatheca length about 1.2 times diame- 
ter of segment, thick-walled ectal duct 
about V2. total length, bulb oval to sacculate, 
ental process absent. 

Etymology. — For the prominent dorso- 
lateral ridges on the body segments. 

Description. — Eleven fixed specimens 
average 1.05 mm in length and range from 
0.90 to 1.23 mm long (Fig. lA). Live, un- 
stressed specimens measure about 1.50 mm 
long. The body is essentially rod-like with 



prominent transverse ridges across the dor- 
sal and lateral surfaces of segments 2 
through 8. All ridges contain supernumer- 
ary muscles (Fig. IC, sm), with the poste- 
rior-most ridge being the largest, and fan- 
like. Segment 1 may occasionally show a 
pronounced transverse ridge. However, this 
is a fixation artifact caused by the head re- 
tracting into the segment. The head is great- 
er in diameter than segment 1 and about 
equal to the posterior attachment disc. 

The peristomium consists of two lips: the 
dorsal usually has four slight rises although 
the margin may appear smooth, while the 
ventral lip has a median emargination. 
There are 16 oral papillae around the 
mouth. The jaws are small, <•/(, head di- 
ameter in width, with sharply pointed teeth 
and a dental formula of 5/4 (Fig. IB). The 
dorsal jaw is kidney-shaped and the ventral 
jaw is quadrangular when seen in dorso- 
ventral aspect. The pharynx has three sulci 
and the median is the largest. 

The male reproductive organs in segment 
6 extend about % of the way dorsad (Fig. 
IC). The subcylindrical, glandular atrium is 
about V2 the diameter of the segment long, 
and about Va wide (short and thick). The 
epithelial cells contain densely packed 
granules, and deferent lobes are absent. The 
prostate gland arises from ectal third of 
glandular atrium, the length and diameter 
are subequal to those of the glandular atri- 
um. The ectal portion of the prostate gland 
contains granular cells and the remainder is 
lined with highly vacuolated, "differentiat- 
ed" gland cells. An ental bulb is absent. 
The muscular atrium length is about V2 that 
of the glandular atrium, and enters the bur- 
sa. The bursa is generally ovoid, and about 
V3 the diameter of the segment in length. 
The penial sheath is formed from the ental 
V3 of the bursa and partially surrounds the 
subspherical penis. Although the penis has 
not been observed in the projected position, 
it is referred to as "protrusible" following 
the functional interpretation made by Holt 
(1986). 

The spermatheca length is about 1.2 



258 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






)fWli 



Fig. 1. Pterodrilus annulatus: A, Lateral view of holotype, scale bar = 100 |xm; B, Oblique lateral and 
dorsal views respectively of the dorsal and ventral jaws, scale bar =10 |xm; C, Lateral view of the reproductive 
systems in segments 5 and 6 in the holotype, scale bar = 20 |xm; D, Lateral view of a curled paratype specimen, 
scale bar = 100 |xm; E and F, Lateral views of segment 5 in two paratype specimens to show the range in size 
of the spermatheca, scale bar = 25 fim. Abbreviations: b, bursa; ga, glandular atrium; ma, muscular atrium; pg, 
prostate gland; sm, supernumerary muscles; spb, spermathecal bulb; spd, spermathecal duct; vd, vas deferens. 



times the diameter of segment 5 (Fig. IC). 
The thick- walled, ectal duct is about % the 
total length of the organ. The bulb is oval, 
frequently with a recognizable narrowing 
ental region, but an ental process is absent. 
Variations. — The transverse segmental 



ridges are less obvious when a specimen is 
fixed in the curled position (Fig. ID). The 
shape of the spermathecal bulb ranged from 
a tubular shape, which happened to be re- 
flexed and contained no spermatozoa (Fig. 
IE), to a sacculate bulb that occupied most 



VOLUME 109, NUMBER 2 



259 



of the dorsal segment and was filled with 
spermatozoa (Fig. IF). All other apparent 
variations could be accounted for by con- 
traction of the specimen due to its state at 
the time of fixation. 

Other branchiobdellidans on the host. — 
Ankyrodrilus legeaus Holt, 1965; Cambar- 
incola chirocephala Ellis, 1919; C. fallax 
Hoffman, 1963; C. philadelphicus (Leidy, 
1851); Oedipodrilus oedipus Holt, 1967; 
Pterodrilus hobbsi Holt, 1968; and Xiron- 
odrilus formosa Ellis, 1918. 

Remarks. — Specimens of P. annulatus 
were observed on the dorsal and lateral sur- 
faces of the carapace, abdomen, and the ce- 
phalothoracic appendages of the host. Usu- 
ally specimens were observed in a cleft or 
by a bunch of setae supporting large pop- 
ulations of diatoms. Pterodrilus annulatus 
were seen to ingest these diatoms and they 
appear to form a significant portion of the 
diet based on the intestinal contents of fixed 
specimens. 

Adjacent to the setal clumps and in the 
clefts of the exoskeleton, P. annulatus was 
often observed feeding alongside speci- 
mens of P. hobbsi. The two species showed 
no aggressive behavior towards one anoth- 
er. 

This manuscript was originally submitted 
for publication in 1993. During the subse- 
quent review period the author obtained a 
list of the branchiobdellidan species depos- 
ited in the National Museum. The collection 
contained a specimen (USNM 99811) with 
the unpublished, and therefore unrecog- 
nized name of Pterodrilus annulatus. This 
specimen, along with three others from two 
different species were mounted on the same 
slide. These branchiobdellidans were sub- 
sequently remounted on separate slides. All 
of these specimen were from lot USNM 
99807 (1-769), "Unidentified Branchiob- 
dellids," a collection removed from Orco- 
nectes neglectus neglectus and Orconectes 
ozarkensis by Perry C. Holt on 7 July 1958, 
obtained at Wildcat Creek adjacent to U.S. 
Route 68, Benton County, Arkansas (Worm 
General Catalog, National Museum of Nat- 



ural History, Smithsonian Institution, 
Washington, DC). Two specimens of P. an- 
nulatus were obtained from this lot and 
mounted for species confirmation (USNM 
169999, USNM 170000). All three bran- 
chiobdellidans were of the same species as 
that already described by the author. To 
avoid future problems, the author withdrew 
his manuscript and changes the name of his 
new species to P. annulatus. 

Pterodrilus annulatus has an external ap- 
pearance and size very similar to Cambar- 
incola serratus Holt, 1981, reported from 
Idaho. However, C. serratus differs from P. 
annulatus in that the former has a dental 
formula of 5/5, the differentiated prostate 
gland arises from the glandular atrium very 
close to the muscular atrium entrance, the 
penis fills over two-thirds of the bursal atri- 
um, and the long spermatheca has a sub- 
spherical terminal bulb. Although these an- 
atomical differences appear obvious, the ef- 
fects of fixation and specimen orientation 
frequently make the differences much more 
subtle. 

Specimens of the crayfish Orconectes 
leptogonopodus and O. palmeri longimanus 
were collected from a site adjacent to U.S. 
Route 270, 2.3 miles east of Mount Ida, 
Montgomery County, Arkansas, by R C. 
Holt on 23 June 1960. Four branchiobdel- 
lidans from this collection were mounted on 
a slide, USNM 100356 (= PCH 1090), and 
identified as Cambarincola serrata by Holt 
[all details come from the Worm General 
Catalogue under USNM 100356]. During 
the present investigation, these four speci- 
mens were remounted on separate slides, 
USNM 100356, USNM 170002, USNM 
170003, USNM 170004, and identified as 
P. annulatus. This correction has main- 
tained the reported distributional separation 
of the western C. serratus and the eastern 
Pterodrilus spp. 

Discussion. — Unlike most of the species 
in the genus, P. annulatus can be grouped 
with P. choritonamus and P. missouriensis, 
as they also lack dorsal segmental append- 
ages. The presence of a "differentiated" 



260 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



prostate gland in P. annulatus separates it 
from P. missouriensis, but provides a char- 
acter in common with the rest of the species 
in the genus. 

Holt (1968) reviewed the species of Pter- 
odrilus and discussed in great detail the 
"Primitive Pterodrilus," "Phylogeny of the 
Genus," and the "Places of Origin and Mi- 
gration." Observations on P. annulatus 
have not resolved or clarified any of the 
concepts addressed by Holt, and so no val- 
ue is seen in discussing these issues here. 

Emmended Species Descriptions 

Pterodrilus cedrus Holt, 1968 
Fig. 2A-B, 3E 

Specimens. — Holotype and five para- 
types (USNM 36464, = PCH 1396), and 
USNM 36465-36468. 

Brief description. — Body terete 1.0 to 1.3 
mm long, low dorsal ridges on segments 2 
through 7 and prominent ridge across seg- 
ment 8 with 5 finger-like projections (Fig. 
3E), supernumerary muscles present; dorsal 
lip smooth; oral papillae present; dental for- 
mula 5/4; male genitalia extend upwards 
just beyond the level of the intestine; glan- 
dular atrium length about V3, segment di- 
ameter; prostate gland ends subterminal to 
glandular atrium, prostate [vacuolated] dif- 
ferentiated, ental bulb absent; muscular atri- 
um length about two-thirds the glandular 
atrium; bursa oval, length about Va segment 
diameter, penial sheath filling ental V2 of 
bursa; spermatheca length about 1.3 times 
segment diameter, ectal duct about V2 
length, bulb tubular to oval, ental process 
absent. 

Variations. — The prominence of the dor- 
sal ridges on segments 2 though 7 depends 
on the degree of contraction and extent of 
curvature of the specimen. The shape of the 
spermathecal bulb varies greatly depending 
on the amount of sperm present in the lu- 
men and the degree of displacement as it 
bends over the intestine (Figs. 2A and B). 

Hosts. — Orconectes placidus (Hagen), 
O. rusticus subspecies, O. juvenilis (Hagen) 



and Cambarus tenebrosus Hay from Holt 
(1968:23). 

Distribution. — Clay County, Tennessee 
from Holt (1968:21). 

Remarks. — The holotype and three para- 
types show five short finger-like projections 
which is contrary to the four reported by 
Holt (1968:21). The length of the muscular 
atrium is usually about % that of the sper- 
miducal gland. However, in the holotype it 
is stretched to be of equal length. The sper- 
matheca has an ectal duct Va to V2 the total 
length of the organ, and not % as drawn by 
Holt (1968:22) in his figure 5a. An exami- 
nation of the paratypes support the revised 
ratio. 

Pterodrilus choritonamus Holt, 1968 
Fig. 31 

The type description reports an ental pro- 
cess on the spermathecal bulb (Holt 1968: 
27). However, an examination of the holo- 
type (USNM 36471, = PCH 1395) and two 
paratype (USNM 35472a,b, = PCH 1395) 
specimens revealed that an ental process is 
absent. The ental end of the elongate sper- 
mathecal bulb is simply bent over the in- 
testine and gives the appearance of a "pro- 
cess". The presence or absence of a sper- 
mathecal ental process is one of the criteria 
used in branchiobdellidan taxonomy (Holt 
1968). 

Pterodrilus hobbsi Holt, 1968 
Fig. 2C, 3F 

Specimens. — Holotype (USNM 36486), 
five paratypes (USNM 36487), 194 other 
specimens (USNM 36488-36508), and 
(SRG 31-35). 

Brief description. — body terete, 1.3 to 
1.7 mm long, supernumerary muscles ab- 
sent, dorsal ridge only across segment 8 
with 5 finger-like projections (Fig. 3F); dor- 
sal lip smooth; male genitalia extends to 
dorsad of segment; glandular atrium length 
about half segment diameter; prostate gland 
ends subterminal to glandular atrium, pros- 
tate [vacuolated] differentiated entirely or 



VOLUME 109, NUMBER 2 



261 






Fig. 2. Pterodrilus cedrus: A, Lateral view of the reproductive organs in segments 5 and 6 in the holotype; 
B, Lateral view of the spermatheca in segment 5 in a paratype, scale bar = 25 iJim. Pterodrilus hobbsi: C, 
Lateral view of the reproductive systems in segments 5 and 6 in the holotype, showing the variation in the 
prostate gland with the double headed arrow, scale bar = 25 ixm. 



ental %, ental bulb absent; muscular atrium 
length subequal to glandular atrium, bursa 
subspherical length Va segment diameter, 
penial sheath filling ental % of bursa; sper- 



matheca length about 0.8 to 1.1 times seg- 
ment diameter, duct about Vz length, bulb 
ovoid, ental process absent. 

Variations. — The dorsal finger-like pro- 



262 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



B 






Fig. 3. Lateral view of the dorsal surface of segments 1 to 8 in nine species of Pterodrilus: A, P. distichus; 
B, P. simondsi; C, P. alcicomus; D, P. mexicanus; E, P. cedrus; F, P. hobbsi; G, P. annulatus; H, P. missour- 
iensis; I, P. choritonamus (A, B, and C were redrawn from Holt, 1968), scale bar = approximately 200 jjim. 
Each figure is arranged so that its respective segment 8, as labelled on the top row, is aligned in a column for 
ease of comparison. Abbreviations: 8, segment 8. 



jections vary in length and spacing, there- 
fore this is a dubious character for separat- 
ing P. hobbsi from P. cedrus. The glandular 
atrium and prostate glands also differ 
slightly in relative length, and the extent of 
vacuolation in the prostate varies (Fig. 3, 
double-headed arrow). The size of the sper- 
mathecal bulb depends upon the amount of 
sperm product being stored and may be as 
small as % the size shown in Fig. 2C. 

Hosts. — Cambarus tenebrosus Hay, C. 
longulus longirostris Faxon, C. parvoculus 
Hobbs & Shoup, C. longulus chasmodac- 
tylus James, C. robustus Girard, C. veter- 
anus Faxon, C. friaufi Hobbs, C. extraneus 
Hagen, C. bartonii cavatus Hay, C. scioten- 



sis Plhoades, C. distans Rhoades, C. bar- 
tonii bartonii (Fabricius), C. longulus lon- 
gulus Girard, C. latimanus (LeConte), C. 
stratus Hay, Cambarus species, C. bartonii 
subspecies; Orconectes erichsonianus (Fax- 
on), O. juvenilis (Hagen), O. rusticus for- 
ceps (Faxon), and Orconectes species — 
from Holt (1968:20) and Orconectes pla- 
cidens (Hagen, 1870). 

Distribution. — This is summarized by 
Holt (1968) in his figure 9 and in a list of 
river systems and counties. For consistency, 
a distribution by counties has been com- 
piled from the Worm General Catalogue of 
the material that Holt (1968:20) deposited 
in the museum: 



VOLUME 109, NUMBER 2 



263 



Lauderdale County, Alabama; Harlan 
County, Kentucky; Alleghany County, 
North Carolina; Anderson, Grainger, Haw- 
kins, Johnson, Lawrence, McMinn, Over- 
ton, Roane, Unicoi, and Union counties, 
Tennessee; Bland, Carroll, Dickenson, Lee, 
Scott, and Washington counties, Virginia, 
and a new record from the Harpeth River, 
Williamson County, Tennessee (see type lo- 
cation of P. annulatus). 

Remarks. — The description and drawings 
given here are based on the holotype, and 
differ from the type description figures giv- 
en by Holt (1968:18). All of the material 
examined is consistent with the illustrations 
given (Figs. 2C, 3F). 

Key to species of Pterodrilus 

(Fig. 3) 

la. dorsal projections on any of segments 

2-5 2 

b. dorsal projections absent from seg- 
ments 2-5 4 

2a. two finger-like projections on segments 

2-7 (Fig. 3 A) P. distichus 

b. fan-like projections on segments 3-5, 

and 8 3 

3a. fan-like dorsal projections on segment 

2 (Fig. 3B) P. simondsi 

b. fan-like dorsal projections absent from 

segment 2 (Fig. 3C) P. alcicornus 

4a. dorsal projections on prominent ridge 

of segment 8 5 

b. dorsal projections absent from promi- 
nent ridge of segment 8 7 

5a. four finger-like projections present (Fig. 

3D) P. mexicanus 

b. five finger-like projections present ... 6 
6a. ridges on segments 2-7 present (Fig. 

3E) P. cedrus 

b. ridges on segments 2-7 absent (Fig. 3F) 

P. hobbsi 

7a. ridges on segments 2-7 high (Fig. 3G) 

P. annulatus 

b. ridges on segments 2-7 low (Fig. 3H) 

P. missouriensis 

c. ridges on segments 2-7 absent (Fig. 31) 

P. choritonamus 



Note: In 6b and 7c where ridges are absent, 
so are the supernumerary muscles. All species 
identifications made using keys should be con- 
firmed with additional anatomical characters. 

Acknowledgments 

My thanks to Drs R. O. Brinkhurst and 
R. D. Kathman for help and hospitality dur- 
ing collecting trips, and the supply of ad- 
ditional crayfish; to Dr. Horton H. Hobbs, 
Jr., for identifying the crayfish; and to Ms. 
Cheryl Bright for help and prompt supply 
of specimens from the National Museum of 
Natural History collection. I am grateful to 
Drs. Kathryn A. Coates and Andrea M. 
Gorman for their constructive criticism of 
the manuscript. 

Literature Cited 

Brinkhurst, R. O., & S. R. Gelder. 1991. Annelida: 
Oligochaeta and Branchiobdellida. Pp. 401-435 
in J. H. Thorpe and A. R Covich, eds.. Ecology 
and classification of North American freshwater 
invertebrates. Academic Press, New York. 

Goodnight, C. J. 1940. The Branchiobdellida (Oli- 
gochaeta) of North American crayfish. — Illinois 
Biological Monographs 17:1-75. 

Hagen, H. A. 1 870. Monograph of the North Ameri- 
can Astacidae. — Illustrated catalogue of the 
Museum of Comparative Zoology at Harvard 
College 3:1-109. 

Holt, P. C. 1968. The genus Pterodrilus (Annelida: 
Branchiobdellida). — Proceedings of the United 
States National Museum 125:1-44. 

. 1981. A resume of the members of the genus 

Cambarincola (Annelida: Branchiobdellida) 
from the Pacific drainage of the United 
States. — Proceedings of the Biological Society 
of Washington 94:675-695. 

. 1986. Newly established families of the order 

Branchiobdellida (Annelida: Clitellata) with a 
synopsis of the genera. — Proceedings of the Bi- 
ological Society of Washington 99:676—702. 

, & B. D. Opell. 1993. A checklist of and il- 



lustrated key to the genera and species of the 
Central and North American Cambarincolidae 
(Clitellata: Branchiobdellida). — Proceedings of 
the Biological Society of Washington 106:251- 
295. 
Moore, J. P. 1895. Pterodrilus, a remarkable discod- 
rilid. — Proceedings of the Academy of Natural 
Sciences of Philadelphia, pp. 449-454. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):264-285. 1996. 

Parkins karenwishnerae, a new genus and species of calanoid copepod 
(Parkiidae, new family) from benthopelagic waters of the eastern 

tropical Pacific Ocean 

Frank D. Ferrari and E. L. Markhaseva 

(FDF) Department of Invertebrate Zoology (MRC 534), National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 20560, U.S.A.; 

(ELM) Academy of Sciences, Zoological Institute, Universitetskaya nab. 1, 

199034 St. Petersburg V-34, Russia 

Abstract. — The last five copepodid stages of a new genus and species of 
calanoid copepod, Parkius karenwishnerae, are described. The basis of the 
maxilliped of P. karenwishnerae is elongate distal to its two medial setae and 
its medial row of denticles; endopodal segments 2-5 are attenuate, forming a 
hook-like structure distally, with denticles along the concave margin. These 
character states are expected to be found in adult females of other species of 
Parkius. Parkiidae, new family, are clausocalanoidean copepods whose adult 
female has two Bradford's setae on the fifth enditic lobe on the basis of maxilla 
2, two medial setae on the basis of the maxilliped, and endopodal segments of 
the maxilliped with 4, 0, 1, 1, 1 setae. On leg 1 the anterior denticles of Von 
Vaupel Klein's organ are found proximal to the presumptive boundary of the 
second endopodal segment. 



Species in four families of calanoid co- 
pepods (Diaixidae Sars, 1902, Phaennidae 
Sars, 1902, Scolecitrichidae Giesbrecht, 
1892, Tharybidae Sars, 1902) possess Brad- 
ford's setae on the terminal part of maxilla 
2 and the syncoxa of the maxilliped. These 
setae are weakly-sclerotized and usually are 
without setules or with setules clustered at 
the tip to form a brush-like structure; they 
often are considered sensory in function 
and were named by Ferrari & Steinberg 
(1993) to recognize the contribution of Dr. 
Janet Bradford to the systematics of these 
families. The four families are presumed to 
constitute a monophyletic group within the 
calanoid copepod superfamily Clausocalan- 
oidea (see Fleminger 1957); we refer to 
them here as bradfordian families. Presently 
there are 283 nominal species among 31 
genera in these families (Table 1). Because 
most of the species are found below the ep- 
ipelagic waters of all oceans, the expecta- 
tion that new genera will be discovered as 



deeper pelagic and benthopelagic waters are 
sampled more thoroughly is fulfilled here 
with a description of a new species of cal- 
anoid copepod with similarly modified se- 
tae but with a distinctively modified max- 
illiped. A new genus and new a new brad- 
fordian family are proposed for this cope- 
pod. Because the morphology of maxilla 1, 
maxilla 2, maxilliped, and endopod of leg 
1 is considered important in establishing re- 
lationships among bradfordian families, 
these appendages are redescribed for Diaix- 
is hibernica (Scott, 1896) and Tharybis ma- 
crophthalma Sars, 1902, which are the type 
species of the type genera of the Diaixidae 
and Tharybidae respectively. 

Methods 

Copepods were collected in the eastern 
tropical Pacific Ocean at Volcano 7 with a 
multiple sampling opening-closing net sys- 
tem with mouth opening of 40 cm by 40 



VOLUME 109, NUMBER 2 



265 



Table 1. — Genera and number of species in five 
bradfordian families. 





No. 




species 


Diaixidae Sars, 1902 




Anawekia Othman & Greenwood, 1994 


1 


Diaixis Sars, 1902 


9 


Parkiidae new family 




Parkius new genus 


1 


Phaennidae Sars, 1902 




Brachycalanus Farran, 1905 


4 


Cephalophanes Sars, 1907 


3 


Cornucalanus Wolfenden, 1905 


8 


Onchocalanus Sars, 1905 


8 


Phaenna Glaus, 1863 


2 


Xantharus Andronov, 1981 


1 


Xanthocalanus Giesbrecht, 1892 


53 


Scolecitrichidae Giesbrecht, 1892 




Amallothrix Sars, 1925 


46 


Archescolecithrix Vyshkvartzeva, 1989 


1 


Heteramalla Sars, 1907 


1 


Landrumius Park, 1983 


2 


Lophothrix Giesbrecht, 1895 


9 


Macandrewella Scott, 1909 


8 


Mixtocalanus Brodsky, 1950 


1 


Parascaphocalanus Brodsky, 1955 


1 


Pseudophaenna Sars, 1902 


1 


Puchinia Vyshkvartzeva, 1989 


1 


Racovitzanus Giesbrecht, 1902 


6 


Scaphocalanus Sars, 1900 


37 


Scolecithricella Sars, 1902 


26 


ScolecithrixBmdy, 1883 


4 


Scolecocalanus Farran, 1936 


3 


Scopalatum Roe, 1975 


6 


Scottocalanus Sars, 1905 


13 


Undinothrix Tanaka, 1961 


1 


Tharybidae Sars, 1902 




Neoscolecithrix Canu, 1896 


4 


Parundinella Fleminger, 1957 


4 


Tharybis Sars, 1902 


9 


Undinella Sars, 1900 


10 



cm and net mesh of 183 microns (Wishner 
& Meise-Munns 1984). During three dives 
(2146-2148) of the submersible ALVIN, 
horizontal tows were taken for one hr each, 
1-5 m above the bottom at the base of Vol- 
cano 7 (13°23'N, 102°27'W) at depths of 
2945-3010 m; samples were fixed at depth 
with gluteraldehyde (Wishner et al. 1995). 
Specimens were preserved later in the lab- 



oratory in 0.5% propylene phenoxytol/4.5% 
propylene glycol/95.0% water. They were 
cleared in steps through 50.0% lactic acid/ 
50.0% water to 100% lactic acid and ex- 
amined with differential interference optics, 
or stained by adding a solution of chlorazol 
black E dissolved in 70.0% ethanol/30.0% 
water and examined with bright-field op- 
tics. 

The second through sixth copepodid 
stages are designated CII to CVI; CVI is 
the adult. Prosome and urosome are Pr and 
Ur respectively. Thoracic and abdominal 
somites are numbered according to their ap- 
pearence during development as interpreted 
from data of Hulsemann (1991). The first 
and oldest thoracic somite bears the max- 
illiped and is fused with the cephalon. The 
youngest is the seventh; among calanoids it 
is the only thoracic somite without an ap- 
pendage. In adult calanoids the seventh is 
the first somite of the urosome, and in adult 
females it is fused to the second abdominal 
somite to form the genital complex. The 
first and oldest abdominal somite is the 
most posterior; it bears the caudal rami. The 
youngest is immediately anterior to the old- 
est, and the remaining abdominal somites 
anteriorly increase in age and decrease in 
numerical designation. 

Cephalic appendages are abbreviated Al 
= antennule; A2 = antenna; Mn = man- 
dible; Mxl = maxillule; Mx2 = maxilla. 
Appendages on thoracic somites are Mxp = 
maxilliped (thoracopod 1); PI -5 = swim- 
ming legs (thoracopods 2-6). The caudal 
ramus is CR. Designations of appendage 
segments follow Huys & Boxshall (1991) 
except for Mx2 and Mxp; exopod = Re; 
endopod = Ri; medial lobe of a segment = 
li, lateral lobe = le. Ramal segments of 
Mx2 are exopodal and the Mxp has at most 
five endopodal segments (Ferrari 1995). 

Ramal segments on thoracopods Mxp 
and PI -5 are numbered by their appearence 
during development (Hulsemann 1991, Fer- 
rari & Ambler 1992, Ferrari 1995) and not 
proximal-to-distal as is the usual case in co- 
pepod descriptions. On the Mxp (Figs. 3F, 



266 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



5D, 8C, J) the distal segment is the first 
endopodal segment, and the second endo- 
podal segment is immediately proximal to 
the first. The third endopodal segment is 
immediately distal to the basis. The fourth 
endopodal segment is immediately distal to 
the third. The fifth endopodal segment is 
the middle segment. The second and first 
segments of a Mxp with a 5-segmented en- 
dopod are more distal and the third and 
fourth segments are more proximal. The 
distal segment of a ramus of Pl-4 (Figs. 
4E, F, 6F, 8E) is the first segment. The sec- 
ond segment is immediately distal to the ba- 
sis. If present, the third segment is imme- 
diately proximal to the distal (or first) seg- 
ment. For a 3-segmented ramus, the proxi- 
mal segment is the second segment, the 
middle segment is the third segment, and 
the distal segment is the first segment. In 
the text, the number of setae recorded for 
ramal segments follows these schemes. 

Armament elements of appendages here 
are termed setae regardless of their position 
or degree of rigidity. Examples of the po- 
sition and morphology of setae are shown 
in the illustrations. Two setae and one 
aesthetasc on a segment of Al are desig- 
nated 2 -I- 1 . Setules are epicuticular exten- 
sions of a seta; denticles are epicuticular ex- 
tensions of an appendage segment; spinules 
are epicuticular extensions of a somite. Von 
Vaupel Klein's organ (Ferrari & Steinberg 
1993, Ferrari 1995) on PI (the appendage 
of thoracic somite 2) consists of the curved 
basal seta, and sensilla, denticles, or pores 
on the anterior face of the endopod. 

Parkiidae, new family 

Diagnosis. — Calanoid copepods with 
adult females having 3 inner setae on the 
first exopodal segment on leg 1 and a 1- 
segmented endopod, and 4 inner setae on 
the first exopodal segment and a 2-seg- 
mented endopod on leg 2. Maxilla 2 with 2 
Bradford's setae on the fifth enditic lobe on 
the basis; with 3 thick, unarmed Bradford's 
setae and 5 with apical setules on the sixth 



enditic lobe plus exopod. Maxilliped with 3 
Bradford's setae, 2 unarmed and 1 with api- 
cal setules, on the syncoxa; with 2 medial 
setae on its basis; with 4, 0, 1, 1, 1 setae 
on the endopod. Von Vaupel Klein's organ 
on leg 1 includes a well-defined group of 
long, thin denticles on the proximal third of 
the anterior face of the endopod; these den- 
ticles insert in an area proximal to the pre- 
sumptive boundary of the second segment. 
Four derived character states of adult fe- 
males exhaustive for the family are (1) 
maxilla 2 with 2 Bradford's setae on the 
fifth enditic lobe on the basis; (2) maxilli- 
ped with 2 medial setae on the basis; (3) 
endopodal segments of the maxilliped with 
4, 0, 1, 1, 1 setae; and (4) leg 1 endopod 
with area of denticles of Von Vaupel 
Klein's organ in proximal position. 

Parkins, new genus 

Diagnosis. — Adult parkiid females (1) 
maxilliped with basis elongate distal to its 
two medial setae and medial denticle row; 
(2) with endopodal segments 2-5 attenuate, 
forming a hook-like structure distally, with 
denticles along the concave margin. 

Type species. — Parkius karenwishnerae, 
by monotypy. 

Etymology. — The genus honors Dr. Tai- 
soo Park for his contributions to the phy- 
logeny and taxonomy of calanoid copepods. 

Parkius karenwishnerae, new species 
Figs. 1-8 

Material examined. — A dissected female 
holotype from dive 2148 at the Zoological 
Institute, St. Petersburg (ZISP 66823). Re- 
maining copepods (CII — 6 specimens; 
CIII — 6 specimens; CIV — 2 females and 4 
males; CV — 2 males; CVI — 2 females) 
comprise one lot of paratypes in the Na- 
tional Museum of Natural History, Smith- 
sonian Institution (USNM 278203). All 
specimens were collected at the base of 
Volcano 7 (13°23'N, 102°27'W), 1-5 m 
above the bottom, at depths of 2945-3010 
m. 



VOLUME 109, NUMBER 2 



267 



CVI female. — Length of 3 specimens 
1.80 (holotype), 2.04, 2.15; average Pr 
length/Ur length = 3.6. 

Pr (Fig. lA): 5 segments; 1st a complex 
of 5 cephalic somites + Thl and 2; Th3-6 
simple and articulated; posterior margin of 
Th6 extended. 

Ur (Fig. lA-C): 4 segments; 1st a genital 
complex of Th7 and Abd2 (Fig. IB, C) 
seminal receptacles curve dorsally and an- 
teriorly; Abd3, 4, 1 articulated. 

Rostrum (Fig. ID): a simple plate with a 
pair of thin filaments; labrum and paragnath 
as illustrated. 

Al (Fig. lE-G): 24 articulated segments 
with 3, 6 + 1, 2 + 1, 2, 2 + 1, 2, 2 + 1, 

4 + 1, 1, 1, 2 + 1, 1, 2 + 1, 2, 2, 2, 2, 1 
+ 1, 1, 1, 2, 2, 2, 4 + 1 setae + aesthetascs. 

A2 (Fig. 2A, B): coxa with 1 seta and a 
row of long denticles; basis with 2 setae. 
Re 6-segmented with 0, 1, 1, 1, 1, 4 setae. 
Ri 2-segmented with 1, 13 (6 terminal, 7 
subterminal) setae. 

Mn (Fig. 2C, D): coxa with 2 areas of 
denticles; basis with 3 setae. Re 5-seg- 
mented with 1, 1, 1, 1, 2 setae. Ri 2-seg- 
mented with 2, 9 setae. 

Mxl (Fig. 3A-C): le with 8 setae. Re 1- 
segmented with 8 setae and several denti- 
cles; baseoendopod with sets of 4, 2, 3, and 

5 setae. Li 2, 3 both with 4 setae. Li 1 with 
9 apical and 2 posterior setae; denticles on 
anterior surface. 

Mx2 (Fig. 2E, F): h 1-4 of coxa each 
with 4, 3, 3, 3 setae each; 115 on basis with 
4 setae, 2 are Bradford's setae without se- 
tules. Li6 + Re an indistinctly segmented 
complex with 8 Bradford's setae; terminal 
3 setae thick without setules, 5 shorter with 
apical setules. 

Mxp (Fig. 3D-F): syncoxa with denticles 
on disto-medial margin and 8 setae (1 short 
Bradford's with apical setules and 2 long 
Bradford's without apical setules). Basis 
elongate with medial row of denticles fol- 
lowed by 2 setae arising from an unscler- 
otized area; indistinct disto-medial lobe 
with 2 setae. Ri 5-segmented with 4, 0, 1, 
1, 1 setae; intersegmental arthrodial mem- 



branes indistinct. Ri 2-5 attenuate, forming 
a hook-like structure with denticles along 
concave margin. 

PI (Fig. 4A, B): coxa with medial den- 
ticles; basis with medial seta and denticles. 
Re 3-segmented with 5 (3 medial, 1 ter- 
minal, 1 lateral), 1 (lateral), 2 (medial and 
lateral) setae; Rel with lateral denticles and 
Re2 and 3 with medial denticles. Ri a 1- 
segmented complex with 5 setae (3 medial, 
2 terminal). Von Vaupel Klein's organ with 
an area of long, thin denticles on proximal 
one-third of anterior face of Ri. A breaking 
plane on most inner setae of Ri; 2 breaking 
planes on most inner setae of Re. 

P2 (Fig. 4C, D): coxa with medial seta 
and denticles. Basis unarmed. Re 3-seg- 
mented with 8 (4 medial, 1 terminal, 3 lat- 
eral), 2 (medial and lateral), 2 (medial and 
lateral) setae; posterior face of Rel and 3 
with distally polarized denticles. Ri 2-seg- 
mented with 5 (2 medial, 2 terminal, 1 lat- 
eral), 1 (medial) setae; posterior face of Ril 
with distally polarized denticles. A break- 
ing plane on most inner setae of Ri; 2 
breaking planes on most inner setae of Re. 

P3 (Fig. 4E, F): coxa with medial seta 
and denticles. Basis unarmed Re 3-seg- 
mented with 8 (4 medial, 1 terminal, 3 lat- 
eral), 2 (medial and lateral), 2 (medial and 
lateral) setae; posterior face of Re 1-3 with 
distally polarized denticles. Ri 3-segmented 
with 5 (2 medial, 2 terminal, 1 lateral), 1 
(medial), 1 (medial) setae; posterior face of 
Ril -3 with distally polarized denticles. A 
breaking plane on most inner setae of Ri; 2 
breaking planes on most inner setae of Re. 

P4 (Fig. 4G, H): coxa with medial seta, 
and medial and posterior denticles. Basis 
with posterior denticles. Re 3-segmented 
with 8 (4 medial, 1 terminal, 3 lateral), 2 
(medial and lateral), 2 (medial and lateral) 
setae; posterior face of Re 1-3 with distally 
polarized denticles. Ri 3-segmented with 5 
(2 medial, 2 terminal, 1 lateral), 1 (medial), 
1 (medial) setae; posterior face of Ril-3 
with distally polarized denticles. A break- 
ing plane on most inner setae of Ri; 2 
breaking planes on most inner setae of Re. 



268 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. I. Parkius karenwishnerae, new genus and species, CVI female: A, animal, left lateral (CR cross- 
hatched); B, genital complex, ventral; C, genital complex, right lateral; D, rostrum, labrum and labium, right 
lateral; E, Al free segments 1-8; F, Al free segments 9-17; G, Al free segments 18-24; H, P5; I, CR. Wavy 
line cutoff indicates broken setae. Line 1 = 0.1 mm (A); line 2 = 0.1 mm (D, E, F, G); line 3 = 0.1 mm (I); 
line 4 = 0.1 mm (B, C); line 5 = 0.1 mm (H). 



VOLUME 109, NUMBER 2 



269 




Fig. 2. Parkius karenwishnerae, new genus and species, CVI female: A, A2, Re; B, A2, coxa, basis and Ri; 
C, Mn, gnathobase, anterior; D, Mn, palp, anterior; E, Mx2, li 1-4 on coxa and li 5 on basis, posterior; F, Mx2, 
li 5 on basis (without setae) and li 6 + Re, posterior Line 1 = 0. 1 mm (A), B; line 2 = 0. 1 mm (E), F; line 3 
= 0.1 mm (C, D). 



270 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 3. Parkius karenwishnerae, new genus and species, CVI female: A, Mxl, syncoxa with li 1 detached, 
posterior; B, Mxl, li 1, posterior; C, Mxl, palp, posterior; D, Mxp, syncoxa; E, Mxp, basis and Ri; F, Mxp, 
detached segments of Ri (b = distal tip of basis; numbers to right indicate relative appearence of ramal segments 
during development). Line 1 = 0.1 mm (F); line 2 = 0.1 mm (A, B, C); line 3 = 0.1 mm (D, E). 



P5 (Fig. IH): coxa and basis without se- 
tae, with posterior denticles. Re 1 -segment- 
ed with 2 setae (medial and lateral); api- 
cally with 2 attenuate points. 



CR (Fig. II): 3 large, terminal setae, 1 
large, postero-lateral seta, 1 small seta on a 
distomedial, ventral lobe, and 1 small seta 
on a dorsal lobe. 



VOLUME 109, NUMBER 2 



271 




Fig. 4. Parkius karenwishnerae, new genus and species, CVI female: A, PI, coxa, basis and Ri, anterior 
(arrow indicates approximate position of presumptive boundary between Ril and Ri2, between proximal two 
medial setae); B, PI, Re, anterior; C, P2, coxa, basis and Ri, posterior; D, P2, Re, posterior; E, P3, coxa, basis 
and Ri, posterior (numbers to left indicate relative appearence of ramal segments during development); F, P3, 
Re (numbers to right indicate relative appearence of ramal segments during development), posterior; G, P4, 
coxa, basis and Ri, posterior; H, P4, Re, posterior. Wavy line cutoff indicates broken setae. Line 1=0.1 mm 
(A, B); line 2 = 0.1 mm (C-H). 



272 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



CVI male. — Not found. 

CV male. — Differs from CVI female as 
follows: length of 2 specimens 1.96, 2.11 
mm; average Pr length/Ur length = 3.5. 

Ur (Fig. 5A): 4 segments; Th7 and Abd 
2, 3, 1 articulated. 

A2 (Fig. 5B): Ri terminal segment with 
12 setae (6 terminal, 6 subterminal). 

Mn (Fig. 5C): Ri2 with 8 setae. 

Mxp (Fig. 5D): Ri 4-segmented with 4, 

0, 1, 1 setae. 

PI: denticles on anterior face of Ri ab- 
sent. 

P5 (Fig. 5E): Re 2-segmented with 1 
(medial), 1 (lateral) setae. Ri 1 -segmented, 
apically attenuate. 

CV female. — Not found. 

CIV female. — Differs from CV male as 
follows: length of 2 specimens 1.37, 1.50 
mm; average Pr length/Ur length = 3.7. 

Pr (Fig. 5F): 6 segments; 1st a complex 
of 5 cephalic somites + Th 1 ; Th2-6 artic- 
ulated; posterior margin of Th6 extended. 

Ur (Fig. 5F): 3 segments; Th7 and Abd2, 
1 articulated. 

Al (Fig. 6A, B): 24 segments; proximal 
8 segments with 3, 3 + 1, 1, 1, 1 + 1, 1, 

1, 2 + 1 setae+aesthetascs. 

Mn (Fig. 5G): Ri2 with 7 setae. 

PI (Fig. 6C): Re 2-segmented with 7 (4 
medial, 1 terminal, 2 lateral), 1 (lateral) se- 
tae. Coxa without medial denticles. 

P2 (Fig. 6D): Re 2-segmented with 9 (5 
medial, 1 terminal, 3 lateral), 2 (medial and 
lateral) setae. Ri with fewer distally polar- 
ized denticles. 

P3 (Fig. 6E): Re 2-segmented with 9 (5 
medial, 1 terminal, 3 lateral), 2 (medial and 
lateral) setae. Ri 2-segmented with 6 (3 me- 
dial, 2 terminal, 1 lateral), 1 (medial) setae; 
posterior face of Ri2 without distally polar- 
ized denticles. 

P4 (Fig. 6F): Re 2-segmented with 9 (5 
medial, 1 terminal, 3 lateral), 1 (lateral) se- 
tae. Ri 2-segmented with 6 (3 medial, 2 ter- 
minal, 1 lateral), 1 (medial) setae. 

P5 (Fig. 5H): Re 1 -segmented with 1 
(lateral) seta. 

CIV male. — Differs from CIV female as 



follows: length of 4 specimens 1.47, 1.48, 
1.53, 1.57 mm; average Pr length/Ur length 
= 3.9. 

Al (Fig. 6A, B): segments 14-16 on 
right appendage each with 2 setae. 

P5 (Fig. 51): Re 1 -segmented with 1 (lat- 
eral) seta. Ri 1 -segmented unarmed. 

C///.— Differs from CIV male as fol- 
lows: length of 6 specimens 1.12, 1.14 (3 
specimens), 1.16, 1.26; average Pr length/ 
Ur length = 3.8. 

Ur (Fig. 7A): 2 segments; Th7 and Abdl 
articulated. 

Al (Fig. 7B— D): 23 articulated segments 
with 3, 1, 1, 1 + 1, 0, 1, 1 + 1, 0, 1, 1, 1, 
1, 1, 1, 1, 1, 1 + 1, 1, 1, 2, 2, 2, 4 + 1 
setae + aesthetascs. 

A2 (Fig. 8A): Ri terminal segment with 
10 setae (5 terminal, 5 subterminal). 

Mn (Fig. 8B): Ri2 with 6 apical setae. 

Mxl (Fig. 7E): Le with 7 setae. Re 1- 
segmented with 6 setae; baseoendopod with 
sets of 3, 3, and 5 setae. Lil with 9 apical 
setae, 1 apical seta reduced in size. 

Mxp (Fig. 8C): disto-medial lobe of basis 
with 1 seta. Ri 4-segmented with 4, 0, 0, 
setae. 

P2: Re denticles absent; sparse on Ri. 

P3 (Fig. 8D): Re 2-segmented with 7 (4 
medial, 1 terminal, 2 lateral), 1 (lateral) se- 
tae; posterior face of Rel without distally 
polarized denticles; Re2 without denticles. 
Ri2 distally polarized denticles sparse. 

P4 (Fig. 8E): coxa unarmed. Re 1 -seg- 
mented with 7 (3 medial, 1 terminal, 3 lat- 
eral) setae. Ri 1 -segmented with 6 (3 me- 
dial, 2 terminal, 1 lateral) setae. 

P5 (Fig. 7F): a simple unarmed lobe on 
medial face of Th6. 

C//.— Differs from ClII as follows: 
length of 6 specimens 0.91 (2 specimens), 
0.93 (2 specimens), 0.95, 1.05 mm; average 
Pr length/Ur length = 3.5. 

Pr (Fig. 8F): 5 segments; Th2-5 articu- 
lated; posterior edge of Th5 in shape of 
simple papilla. 

Ur (Fig. 8F): 2 segments; Th6 with lat- 
eral lobes and Abdl articulated. 

Al (Fig. 7G, H): 19 articulated segments 



VOLUME 109, NUMBER 2 



273 




Fig. 5. Parkius karenwishnerae, new genus and species, CV male: A, animal, right lateral (CR cross- 
hatched); B, A2, Ri2; C, Mn, Ri2; D, Mxp, Ri (b = distal tip of basis; numbers below indicate relative appearence 
of ramal segments during development); E, P5. CIV female: F, animal, left lateral (CR cross-hatched); G, Mn 
Ri2; H, P5. CIV male: I, P5. Line 1 = 0.1 mm (A); line 2 = 0.1 mm (F); line 3 = 0.1 mm (B-E, G-I). 



with 3, 2 + 1, 0, 1, 0, 1, 0, 1, 0, 1, 0, 1, 1, 
0, 1, 2, 2, 2, 4 + 1 setae + aesthetascs. 

A2 (Fig. 8G): Ri terminal segment with 
9 (5 terminal, 4 subterminal) setae. 



Mn (Fig. 8H): Ri2 with 5 setae. 
Mxl (Fig. 71): Le with 6 setae. Li 2 with 
3 setae. Lil with 7 apical setae. 

Mxp (Fig. 81, J): syncoxa with 7 setae 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 6. Parkius karenwishnerae, new genus and species, CFV female: A, schematic of Al, free segments 1- 
10; B, Al, free segments 11-18 (# indicate setae which are present only on right Al of CIV male); C, PI, Re, 
anterior; D, P2, Re, posterior; E, P3, Re and Ri, posterior; F, P4, Re and Ri, posterior (numbers to right of Re 
and left of Ri indicate relative appearence of ramal segments during development). Line 1 = 0.1 mm (A, B); 
line 2 = 0.1 mm (C, E, F); line 3 = 0.1 mm (D). 



VOLUME 109, NUMBER 2 



275 




Fig. 7. Parkius karenwishnerae, new genus and species, CIII: A, animal, right lateral (CR cross-hatched); 
B, Al, free segments 1-14; C, Al, free segments 15-19; D, free segments 20-23; E, Mxl, posterior; F, P5 (tip 
of arrow) near posterior edge of Th6 and Th7 (#). CII: G, Al, free segments 1-12; H, Al, free segments 13- 
19; I, Mxl, posterior; J, P4 at posterior edge of Th5 (Th6 with #). Wavy line cutoff indicates broken setae. Line 
1 = 0.1 mm (A); line 2 = 0.1 rrmi (B-D, G, H); line 3 = 0.1 mm (E, F, I, J). 



276 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 8. Parkins karenwishnerae, new genus and species, CIII: A, A2, Ri2; B, Mn, Ri2; C, Mxp, Ri (b = 
distal tip of basis; numbers to left indicate relative appearence of ramal segments during development); D, P3, 
Re, posterior; E, P4, posterior (numbers to right of Re and left of Ri indicate relative appearence of ramal 
segments during development). CII: F, animal, left lateral (CR cross-hatched); G, A2, Ri2; H, Mn, Ri2; I, Mxp, 
syncoxa; J, Mxp, Ri (b = distal tip of basis; numbers to left indicate relative appearence of ramal segments 
during development); K, P2, Re, posterior; L, P3, posterior. Wavy line cutoff indicates broken setae. Line 1 = 
0.1 mm (F); line 2 = 0.1 mm (D, E, \, K, L); line 3 = 0.1 mm (A-D, H, J). 



VOLUME 109, NUMBER 2 



277 



(only 2 in disto-medial set); Ri 3-segmented 
with 4, 0, setae. 

P2 (Fig. 8K): Re 2-segmented with 7 (4 
medial, 1 terminal, 2 lateral), 1 (lateral) se- 
tae. 

P3 (Fig. 8L): coxa unarmed. Re 1 -seg- 
mented with 7 (3 medial, 1 terminal, 3 lat- 
eral) setae. Ri 1 -segmented with 6 (3 me- 
dial, 2 terminal, 1 lateral) setae. 

P4 (Fig. 7J): a bilobe bud on posterior 
edge of Th5; dorsal lobe with 3 weakly- 
sclerotized setae, ventral lobe with 2 weak- 
ly-sclerotized setae; lobes and setae direct- 
ed dorsally. 

Etymology. — This new species honors 
Dr. Karen Wishner for her many and varied 
contributions to the biology of the oceans. 

Remarks. — Adult females of P. karen- 
wishnerae share with species of the calan- 
oid superfamilies Ryocalanoidea, Spinoca- 
lanoidea, and Clausocalanoidea, a 1 -seg- 
mented endopod on leg 1, and a 2-seg- 
mented endopod on leg 2. Adult males of 
species of the latter two superfamilies have 
non-geniculate antenna 1, a character state 
we have been unable to determine for the 
new species. Adult females of P. karen- 
wishnerae share with species of the Clau- 
socalanoidea three inner setae on the first 
exopodal segment of leg 1 and four inner 
setae on the first exopodal segment of leg 
2 (Park 1986). Adult females of Parkius 
share: with diaixids, tharybids, phaennids 
and scolecitrichids, Bradford's setae on the 
sixth enditic lobe on the basis plus exopod 
of maxilla 2; with diaixids and scolecitri- 
chids three apical Bradford's setae without 
setules and five more with apical setules on 
this segment complex; with diaixids, phaen- 
nids and scolecitrichids distally and/or ra- 
dially polarized denticles on the posterior 
surfaces of some ramal segments of legs 2- 
4; with most scolecitrichids two Bradford's 
setae on the fifth enditic lobe on the basis 
of maxilla 2. The combination of the fol- 
lowing derived character states separates 
Parkiidae from the other bradfordian fami- 
lies: maxilla 2 with two Bradford's setae on 
the fifth enditic lobe on the basis; maxilli- 



ped with two medial setae on the basis; and 
4, 0, 1 , 1 , 1 setae on its endopodal segments 
(reduced by truncation during develop- 
ment); and one leg 1 anterior denticles of 
Von Vaupel Klein's organ proximal to the 
presumptive boundary of the second endo- 
podal segment. 

The shapes of maxillipedal segments are 
diagnostic for Parkius; an elongate basis 
distal to the medial row of denticles and 
distal to the two medial setae; endopodal 
segments 2-5 of the maxilliped attenuate, 
forming a hook-like structure distally, with 
denticles along the concave margin. 

Neither homologies of Bradford's setae 
at various positions on maxilla 2, nor the 
transformation sequence of those setae have 
been hypothesized. We do not know the an- 
cestral state for numbers of Bradford's setae 
on the sixth enditic lobe of the basis plus 
exopod of maxilla 2 although presence of 
one sclerotized seta on Tharybis macro- 
phthalma may be a primitive condition. We 
believe that the presence of four sclerotized 
setae on the fifth enditic lobe on the basis 
is primitive, and that two Bradford's setae 
on that lobe is a derived character state. 
Phylogenetic hypotheses about species 
within bradfordian families await careful 
collecting, better preservation and complete 
descriptions of appendage armament. Of 
particular value to our understanding of re- 
lationships among the species are the ar- 
mament of maxilla 1, presence of Brad- 
ford's setae on the fifth and sixth enditic 
lobes and exopod of maxilla 2, shape and 
armament of maxilla 2, armament of the 
maxilliped, structure of Von Vaupel Klein's 
organ, presence of radially polarized den- 
ticles on legs 2-4, homologies of segments 
and setae of leg 5. 

Segmental and setal homologies. — The 
number of segments and associated setae on 
the ramus of maxilla 2 of the ancestor to 
the bradfordian families is difficult to infer 
based on our present knowledge of clau- 
socalanoideans. Presumed older calanoids 
like Pleuromamma xiphias or Ridgewayia 
klausruetzleri with two enditic lobes on the 



278 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



basis (the fifth and sixth enditic lobes) have 
seven exopodal setae (Ferrari 1985, Ferrari 
1995). Presence of eight setae on the distal 
segments of many species in the bradfor- 
dian families apparently is correlated with 
reduction in size of the sixth enditic lobe. 
The hypothesis accepted here is that the 
sixth lobe, apparently with at least one of 
its setae, are included in the exopod and 
group of eight Bradford's setae, although 
the homologies of each of these setae can- 
not as yet be determined. The tip of maxilla 
2 then is a complex of exopodal segments 
plus the sixth enditic lobe on the basis 
whose setation is complete early in the co- 
pepodid phase of calanoid development. 
This hypothesis is supported by the fact 
that, in general, setation of enditic lobes and 
exopodal segments are complete early in 
calanoid copepodid development (Ferrari 
1995). 

A medial row of denticles is found on 
the proximal portion of the basis of the 
maxilliped of many copepods. Usually this 
row ends at the level of the second of three 
medial setae. On P. karenwishnerae these 
denticles and only two medial setae, occupy 
the proximal one-fourth of the basis; the 
distal portion of the basis is elongate. Some 
species of the phaennid genus Onchocalan- 
us have a maxilliped with an elongate basis 
(Park 1983). However, in O. trigoniceps, O. 
paratrigoniceps, O. magnus, and O. cris- 
tatus the area of elongation is not homolo- 
gous to P. karenwishnerae because it ap- 
parently results from the proliferation of 
cells immediately distal to the medial den- 
ticle row so that cells producing the three 
medial setae of the basis are included in the 
area of elongation. In Parkius karenwish- 
nerae this area of elongation apparently re- 
sults from the proliferation of cells distal to 
the two medial setae as well as the medial 
denticle row, so that cells producing the two 
medial setae are not included in the area of 
elongation. 

Table 2 compares setation during cope- 
podid development of the maxilliped of P. 
karenwishnerae, the related scolecitrichid 



Table 2. — Setation of the maxilliped of Parkius kar- 
enwishnerae and Scopalatum vorax for stages CII-CVI 
and of Ridgewayia klausruetzleri for stages CI-CVI. 
Beginning with CII, setation of the endopod of the 
scolecitrichid, S. vorax, is identical to the presumed 
older calanoid, R. klausruetzleri. P. karenwishnerae 
exhibits delayed appearance of the fifth endopodal seg- 
ment, no seta at the formation of endopodal segments 
2—^, and truncation of post-formation setal addition on 
three endopodal segments. Lobes of the syncoxa (sl- 
s4), the basis (b) and its distomedial lobe (1), and the 
endopodal segments (nl-n5 numbered by develop- 
mental stage) are arranged from left, proximally, to 
right, distally. a = segment not formed. 









Parkius karenwishnerae 












si 


s2 


s3 


s4 


b 


1 


n3 


n4 


n5 


n2 


nl 


CI 


? 


7 


7 


7 


7 


7 


? 


9 


? 


? 


? 


CII 


I 


2 


2 


3 


2 


1 





a 


a 





4 


cm 


1 


2 


2 


3 


2 


1 








a 





4 


CIV 


1 


2 


2 


3 


2 


2 


1 


1 


a 





4 


cv 


1 


2 


2 


3 


2 


2 


1 


1 


a 





4 


cvi 


1 


2 


2 


3 


2 


2 


1 


1 


1 





4 










Scopalatum 


vorax 












si 


s2 


s3 


s4 


b 


1 


n3 


n4 


n5 


n2 


nl 


CI 


7 


7 




7 


7 


7 


? 


? 


9 


9 


9 


CII 


1 


2 




3 


2 


2 


1 


a 


a 


1 


4 


cm 


1 


2 




3 


3 


2 


1 


1 


a 


2 


4 


CIV 


1 


2 




3 


3 


2 


2 


2 


1 


2 


4 


cv 


1 


2 




3 


3 


2 


3 


3 


2 


3 


4 


CVI 


1 


2 




3 


3 


2 


4 


4 


3 


4 


4 








Ridgewayia klausruetzleri 












si 


s2 


s3 


s4 


b 


1 


n3 


n4 


n5 


n2 


nl 


CI 





1 


2 


2 


2 


1 


a 


a 


a 


1 


4 


CII 


1 


2 


4 


3 


3 


2 


1 


a 


a 


1 


4 


cm 


1 


2 


4 


3 


3 


2 


1 


1 


a 


2 


4 


CIV 


1 


2 


4 


3 


3 


2 


2 


2 


1 


2 


4 


cv 


1 


2 


4 


3 


3 


2 


3 


3 


2 


3 


4 


CVI 


1 


2 


4 


3 


3 


2 


4 


4 


3 


4 


4 



Scopalatum. vorax, and R. klausruetzleri, a 
presumed older calanoid. Ferrari & Stein- 
berg (1993) incorrectly figured and de- 
scribed S. vorax with three setae on the 
proximal (third) endopodal segment at CIV; 
the correct number for that stage is two. 
There are no differences between S. vorax 
and R. klausruetzleri in segmentation or se- 
tation of the endopod. However, S. vorax 
has three fewer setae on a lobe of the syn- 
coxa and the addition to the basis of the 
third medial seta is delayed until CIII. Par- 
kius karenwishnerae has two fewer setae on 



VOLUME 109, NUMBER 2 



279 



the homologous syncoxal lobe, a third me- 
dial seta is never added to the basis, the 
addition of the second seta to the disto-me- 
dial lobe of the basis is delayed until CIV, 
and the addition of the fifth endopodal seg- 
ment is delayed until CVI. These changes 
are neotenic because appearances of seg- 
ments or setae are delayed until late in de- 
velopment or delayed so that they do not 
appear at all. There is no seta present at the 
formation of endopodal segments 2-4 of P. 
karenwishnerae, and only one of three plus 
three post-formation setae are added to en- 
dopodal segments 3 and 4. The pattern of 
endopodal setation exhibited by R. klaus- 
ruetzleri and S. vorax appears to have been 
truncated in P. karenwishnerae, resulting in 
a reduction in setal number on the endopod. 
An alternate hypothesis of neotenic reduc- 
tion, i.e., delaying setal addition, is not sup- 
ported by the pattern for P. karenwishnerae 
because final setal numbers are reached at 
CIV rather than CVI. Thus two develop- 
mental processes, neoteny and truncation, 
may be responsible for setal addition to the 
maxilliped of P. karenwishnerae. 

The morphology of Von Vaupel Klein's 
organ suggests that the 1 -segmented endo- 
pod of leg 1 of P. karenwishnerae is a seg- 
ment complex. The development of leg 1 
of P. karenwishnerae is incompletely re- 
ported here, but we assume that its mor- 
phology at CI is similar to Drepanopus for- 
cipatus with 1 -segmented exopod and en- 
dopod (Hulsemann 1991), and without pre- 
sumptive structures of Von Vaupel Klein's 
organ (also see Ferrari, 1995, for develop- 
ment of R. klausruetzleri). On D. forcipa- 
tus, S. vorax or P. karenwishnerae this en- 
dopod does not add segments or setae later 
in development; however, structural ele- 
ments of Von Vaupel Klein's organ (tuber- 
cle and curved basal seta) presumabley are 
added during the molt to CII. At this stage 
in development of presumed older calan- 
oids like R. klausruetzleri, which continue 
to add segments and setae to leg 1 endopod 
during development, the endopod is 2-seg- 
mented. For CII of R. klausruetzleri the 



boundary between the second (proximal) 
and first (distal) endopodal segments of a 
2-segmented endopod has formed between 
the 2 medial setae of the endopod of CI, 
and the tubercule of Von Vaupel Klein's or- 
gan is located near that boundary. Later in 
development the tubercule is located near 
the boundary of the second (proximal) and 
third (middle) endopodal segments of the 3- 
segmented endopod at CV of R. klausruet- 
zleri and P. xiphias. Based on the position 
between the two proximal inner setae of 
this presumed homologous denticule bear- 
ing tubercule, we believe that the 1 -seg- 
mented endopod of leg 1 of D. forcipatus, 
species of Euchirella and Pseudochirella 
mentioned by Von Vaupel Klein (1972), 
and S. vorax is a complex of cells of the 
first and second endopodal segments which 
are not separated by an arthrodial mem- 
brane. The 1 -segmented ramus of P. kar- 
enwishnerae also is a segment complex, but 
without a distinct tubercle. The homologous 
denticles are located away from the bound- 
ary of the presumed second and first seg- 
ments. The alternate hypothesis, that the 1- 
segmented endopod of leg 1 is simply the 
first endopodal segment of CI, would re- 
quire Von Vaupel Klein's organ to have 
evolved independently among calanoids 
with 1 -segmented and with 2- or 3-seg- 
mented endopods on leg 1. 

The pattern of development of legs 1-4 
of P. karenwishnerae is identical to S. vor- 
ax although Ferrari & Steinberg (1993) in- 
correctly described the second exopodal 
segments of leg 4 of CIV male and of leg 
3 of CIII with two setae (they have one 
seta), and the endopod of leg 4 which at 
CIII was incorrectly described and illustrat- 
ed with five setae (there are six; the proxi- 
mal, medial one was not shown). The ramus 
on female leg 5 appears to be an exopod, 
because it is morphologically similar to the 
outer ramus of the male. Distal structures 
on the exopod of female leg 5 exopod of 
P. karenwishnerae and both rami of the 
male do not articulate with the segment but 



280 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



instead appear to be attenuations of the seg- 
ment. 

Bradford (1973) and Bradford et al. 
(1983) have made significant contributions 
to our understanding of the specialized 
clausocalanoidean families Phaennidae and 
Scolecitrichidae despite the fact that the 
morphology of many described species is 
incompletely known. Most of the species 
live in deep water and thus few specimens 
have been available for study; adult males 
are usually rare. In addition, the ramal seg- 
ments of legs 1-4 are often broken during 
capture, and Bradford's setae, which are 
weakly-sclerotized, may become distorted 
over time in ethanol, a commonly used pre- 
servative. A better understanding of rela- 
tionships among bradfordian families has 
been hampered by incomplete descriptions 
of species in the families Diaixidae and 
Tharybidae. Here maxilla 1, maxilla 2, 
maxilliped and endopod of leg 1 are rede- 
scribed for Diaixis hibernica (Scott 1896) 
and Tharybis macrophthalma Sars 1902, 
which are the type species of the type gen- 
era of their respective families. 

Diaixidae Sars, 1902 

Diaixis Sars, 1902 

Diaixis hibernica (Scott 1896) 

Material examined. — CVI 5 females and 
2 males plus several other copepodids from 
Raunefjord, Norway (60°16'N, 5°10'E) col- 
lected by Audun Fosshagen on 4 April 
1995 with a Beyer epibenthic sampler (180 
micron mesh) at 1 20 m have been deposited 
in the U.S. National Museum (USNM 
278204). 

CVI female.— Mx\ (Fig. 9A) Le with 8 
setae. Re 1 -segmented with 8 setae; bas- 
eoendopod with sets of 3, 3, 3, and 3 setae. 
Li2 with 2 setae, Li3 with 3. Li 1 with 8 
apical setae. 

Mx2 (Fig. 9B, C): Li 1-4 on coxa each 
with 4, 3, 3, 3 setae each; 115 on basis with 
4 setae, 1 weakly-sclerotized without set- 
ules. Li2 + Re an indistinctly segmented 
complex with 8 Bradford's setae; terminal 



3 thick without setules, 5 thinner with api- 
cal setules. 

Mxp (Fig. 9D-E): syncoxa with denticles 
on disto-medial margin and 9 setae (1 Brad- 
ford's with setules along its length and 2 
Bradford's with narrow base and without 
apical setules). Basis with medial row of 
denticles and 3 medial setae; disto-medial 
lobe with 2 setae. Ri 5-segmented with 4, 
4, 4, 4, 3 setae. 

PI (Fig. 9F, G): Von Vaupel Klein's or- 
gan with an area of long, thin denticles lat- 
erally on anterior face of Ri. There are also 
3 areas on denticles on the posterior face of 
Ri. 

CVI male. — Differs from CVI female as 
follows. Mxl (Fig. lOA) Le with 4 setae. 
Re 1 -segmented with 7 setae; Ri with 1 me- 
dial and 4 apical setae. Li 1-3 unarmed. 

Mx2 (Fig. lOB): Li 1-3 on coxa with nu- 
merous small denticles; li 4 with a sclero- 
tized seta and denticles; 115 on basis with 1 
Bradford's seta without setules. Li2 with 1 
Bradford's seta with setules; Re indistinctly 
segmented with 5 Bradford's setae; terminal 
3 thick without setules, and 2 shorter with 
apical setules. 

Mxp (Fig. IOC): syncoxa with 4 setae (1 
Bradford's with apical setules). Basis with 
medial row of denticles and 2 medial setae. 

Remarks. — We examined from The Nat- 
ural History Museum in London one lot 
(B.M. 1911.11.8 37900-907) of Scolecith- 
rix hibernica labelled types by A. Scott and 
"Feby 1896" from the Irish Sea which con- 
tained CVI 4 females, 5 males; CV 1 fe- 
male, 2 males, and one lot from the Zoo- 
logical Museum of the University of Oslo 
(F 20677) labeled "Diaixis hibernica A. 
Scott, sted. Bundefj., dt. G. O. Sars" which 
contained CVI 12 females, 1 male; CV 5 
females, 2 males. We did not dissect spec- 
imens from either lot, but adult females 
from both lots agree with Raunefjord spec- 
imens in morphology of the posterior mar- 
gin of the prosome and of the genital com- 
plex; we believe all specimens are conspe- 
cific. For adult females of both lots, we 
were able to verify the general shape of 



VOLUME 109, NUMBER 2 



281 




Fig. 9. Diaixis hibernica (Scott, 1896), female: A, Mxl, posterior; B, Mx2, Iil-4 on syncoxa, posterior (115 
with one seta with circular cutoff also shown); C, Mx2, 115 on basis and 116 + Re, anterior; D, Mxp (setation 
of Ri not shown); E, Mxp, Ri (b = distal tip of basis; numbers to left indicate relative appearence of ramal 
segments during development); F. PI, Ri, anterior (arrow indicates approximate position of presumptive boundary 
between Ril and Ri2, between proximal two medial setae); G, PI, Ri, posterior. Line 1 = 0.1 mm (A-D); line 
2 = 0.1 mm (E); line 3 = 0.1 mm (F, G). 



282 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 10. Diaixis hibemica (Scott, 1896), male: A, Mxl, posterior; B, Mx2; C, Mxp, coxa and basis. Line 1 
= 0.1 mm (A, B); line 2 = 0.1 mm (C). 



VOLUME 109, NUMBER 2 



28a 




Fig. 11. Tharybis macrophthalma Sars, 1902: A, Mxl, anterior; B, Mxl, tip of HI, posterior; C, Mx2, lil- 
4 on syncoxa, posterior; D, Mx2, li5 on basis and 116 + Re, anterior; E, Mxp, syncoxa; F, basis and Ri (setation 
of Ri not shown); G, Mxp, Ri (b = distal tip of basis; numbers to right indicate relative appearence of ramal 
segments during development); H. PI, Ri, anterior (arrow indicates approximate position of presumptive bound- 
ary between Ril and Ri2, between proximal two medial setae). Line 1 = 0. 1 mm (E, F); line 2 = 0.1 mm (A, 
B); line 3 = 0.1 mm (C, D, G); line 4 = 0.1 mm (H). 



284 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



Mxl and setation of the exite, the armament 
of H6 plus Re of Mx2, and setation of basis 
and presence of 19 setae on Ri of Mxp 
which agree with Raunefjord specimens. 

Tharybidae Sars, 1902 

Tharybis Sars, 1902 

Tharybis macrophthalma Sars, 1902 

Material examined. — CVI 9 females and 
10 males from Raunefjord, Norway 
(60°16'N, 5°10'E) collected by Audun 
Fosshagen on 4 April 1995 with a Beyer 
epibenthic sampler (180 micron mesh) at 
120 m have been deposited in the U.S. Na- 
tional Museum (USNM 278205). 

CVI female.— Mx\ (Fig. 11 A, B): Le 
with 9 setae. Re 1 -segmented with 5 setae; 
baseoendopod with sets of 3, 2, 2, and 3 
setae. Li 2 with 3 setae, Li3 with 4. Li 1 
with 9 apical (5 thick with setules, 4 thin 
and unarmed) and 4 posterior setae. 

Mx2 (Fig. lie, D): Li 1-4 of coxa each 
with 4, 3, 3, 3 setae each; li5 on basis with 
4 sclerotized setae. Li2 + Re an indistinctly 
segmented complex with 8 setae; terminal 
3 Bradford's setae without setules, 4 Brad- 
ford's setae with apical setules, and 1 scler- 
otized seta with setules. 

Mxp (Fig. IIE-G): syncoxa with denti- 
cles on disto-medial margin and 9 setae (1 
short Bradford's with apical setules). Basis 
elongate with medial row of denticles fol- 
lowed by 3 setae; disto-medial lobe with 2 
setae. Ri 5-segmented with 4, 4, 4, 4, 3 se- 
tae. 

PI (Fig. IIH): Von Vaupel Klein's organ 
with lateral margin of Ri extended distally; 
denticles medial and lateral to the exten- 
sion, and on the anterior face of Ri below 
the basal seta. 

CVI male. — Mxl, Mx2, Mxp, and PI 
similar to female. 

Remarks. — We examined one lot from 
the Zoological Museum of the University 
of Oslo (F 20610) labeled ''Tharybis ma- 
crophthalma G. O. Sars, sted. Drobak, dt. 
G. O. Sars" which contained CVI 5 fe- 
males, 2 males; CV 1 male. We did not dis- 



sect these specimens but adult females 
agree with Raunefjord specimens in mor- 
phology of leg 5 and the genital complex; 
we believe specimens from both areas are 
conspecific. For adult females we were able 
to verify the armament of 116 plus Re of 
Mx2, and setation of coxa and basis, and 
the presence of 1 9 setae on Ri of Mxp. 

Acknowledgments 

Our thanks to Karen Wishner, University 
of Rhode Island, for the copepods collected 
from the area of Volcano 7, to Audun Foss- 
hagen, University of Bergen, for Tharybis 
macrophthalma and Diaixis hibernica from 
Raunefjord, to Marit E. Christiansen, Uni- 
versity of Oslo, for types of Tharybis ma- 
crophthalma and Sars' specimens of Diaix- 
is hibernica, and to Geoff Boxshall and 
Ann Morgan, The Natural History Muse- 
um, London, for types of Diaixis hibernica. 

Literature Cited 

Bradford, J. 1973. Revision of family and some ge- 
neric definitions in the Phaennidae and Scole- 
cithricidae (Copepoda: Calanoida). — New Zea- 
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search 7:133-152. 

, L. Haakonssen, & J. Jillett. 1983. The marine 

fauna of New Zealand: pelagic calanoid cope- 
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cithricidae, Diaixidae, and Tharybidae. — New 
Zealand Oceanographic Institute Memoir 90, 
150 pp. 

Ferrari, F 1985. Postnaupliar development of a look- 
ing-glass copepod, Pleuromamma xiphias 
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. 1995. Six copepodid stages of Ridgewayia 

klausruetzleri, a new species of copepod crus- 
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barrier reef in Belize, with comments on ap- 
pendage development. — Proceedings of the Bi- 
ological Society of Washington 108:180-200. 

, & J. Ambler. 1992. Nauplii and copepodids 

of the cyclopoid copepod Dioithona oculata 
(Farran, 1913) (Oithonidae) from a mangrove 
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, & D. Steinberg. 1993. Scopalatum vorax (Es- 

terly, 1911) and Scolecithricella lobophora 
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associated with a pelagic tunicate in Monterey 
Bay. — Proceedings of the Biological Society of 
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Fleminger, A. 1957. New genus and two new species 
of Tharybidae (Copepoda Calanoida) from the 
Gulf of Mexico with remarks on the status of 
the family. — Fishery Bulletin 116:347-354. 

Giesbrecht, W. 1892. Systematik und Faunistik den 
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Fauna und Flora des Golfes von Neapel und der 
angrenzenden Meeres-abschnitte 19:1-831 + 
54 pi. 

Hulsemann, K. 1991. The copepodid stages of Dre- 
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the genus and a comparison to other members 
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anoida). — Helgolander Meeresuntersuchungen 
45:199-224. 

Huys, R., & G. Boxshall. 1991. Copepod evolution. 
The Ray Society, London, vol. 159, 468 pp. 

Park, T. 1983. Calanoid copepods of the family 
Phaennidae from antarctic and subantarctic wa- 
ters. — Antarctic Research Series 39:317-368. 



. 1986. Phylogeny of calanoid copepods. — 

Syllogeus 58:191-196. 

Sars, G. 1902. Copepoda Calanoida, part V & VI Sco- 
lecithricidae, Diaixiidae, Stephidae, Tharybi- 
dae, Pseudocyclopiidae. Pp. 49-72 -I- pi. 34-48 
in An account of the Crustacea of Norway, with 
short descriptions and figures of all of the spe- 
cies. The Bergen Museum, Bergen, 4:144 pp + 
96 pi. 

Scott, A. 1 896. On Scolecithrix hibemica, a new spe- 
cies of copepod, with some remarks on the dis- 
tribution of the Crustacea. — Annals and Maga- 
zine of Natural History, series 6, 18:362-367, 
pi. 17-18. 

Vaupel Klein, J. von. 1972. A new character with sys- 
tematic value in Euchirella (Copepoda, Calan- 
oida). — Zoologische Mededelingen 47:497-512 
+ 6 pi. 

Wishner, K., & C. Meise-Munns. 1984. In situ grazing 
rates of deep-sea benthic boundary-layer zoo- 
plankton. — Marine Biology 84:65-74. 

, C. Ashjian, C. Gelfman, M. Gowing, L. Kann, 

L. Levin, L. Mullineaux, & J. Saltzman. 1995. 
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face of the Eastern Tropical Pacific oxygen min- 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2): 286-298. 1996. 



Occurrence of anomuran crabs (Crustacea: Decapoda) in 
hydrothermal vent and cold-seep communities: a review 

Pierre Chevaldonne and Karine Olu 

(PC & KO) Laboratoire d'Ecologie Abyssale, IFREMER, 

B.P. 70, 29280 Plouzane, France; 

(PC) Present address: Institute of Marine and Coastal Sciences, Rutgers University, 

Dudley Road, Cook College, New Brunswick, New Jersey 08903, U.S.A. 

Abstract. — Crabs of the family Lithodidae are frequently encountered in the 
vicinity of deep-sea hydrothermal vents and cold-seeps. Together with crabs of 
the families Galatheidae and Chirostylidae, they are the main contributors to 
the scavenging/predatory fauna of these highly productive areas, and a potential 
vector for the export of organic carbon to the surrounding deep-sea commu- 
nities. A review of the literature indicates that anomuran crabs have been re- 
ported from such environments since their discovery, and occur virtually any- 
where a reducing habitat is found. These three families are represented by at 
least eight genera, with at least 14 species occurring in hot vent areas, and 
eight in cold-seep associated communities. 



Two species of lithodid crabs have re- 
cently been reported from the south Bar- 
bados accretionary prism cold-seeps at 
depths of 1200-1700 m (Macpherson 
1994). In addition to the description of a 
new species, Paralomis arethusa, Macpher- 
son mentioned that his new species and Li- 
thodes manningi Macpherson, 1988, were 
the first records of the family Lithodidae to 
be found in hydrothermal vent and cold- 
seep areas. However, a review of the liter- 
ature showed that occurrence of lithodid 
crabs in such environments has been known 
at least since 1985 (Suess et al. 1985). The 
taxonomic literature on deep-sea hydrother- 
mal vent and cold-seep organisms often 
lacks ecological data, and ecological sur- 
veys frequently present vague taxonomic 
information. This review is intended to 
demonstrate that the Anomura are a partic- 
ularly well represented group in deep-sea 
chemosynthetically-based ecosystems, and 
to present a literature survey of the species 
and accompanying information relevant to 
biogeographic studies. 



Vents and seeps are environments where 
biomass and biological production are high 
compared to that of the surrounding abyssal 
plains, due to utilization of the expelled re- 
duced compounds by chemoautotrophic mi- 
croorganisms that constitute the first step of 
a trophic web independent from sea-surface 
production. Although organisms exclusive- 
ly dependent on fluid emissions usually 
dominate these communities and live close 
to the expelled fluids, other inhabitants of 
the neighboring deep-sea environment are 
attracted by the high food supply and enter 
these environments. Like anomurans, most 
of these organisms are filter-feeders, scav- 
engers, or predators. As top predators/scav- 
engers, anomurans are rivalled only in some 
cases, in particular by the hydrothermal-en- 
demic brachyuran crab family Bythograei- 
dae. 

Infraorder Anomura 

Superfamily Paguroidea (sensu Forest 
1987) 

This superfamily comprises the Lithodi- 
dae and two families of hermit crabs. Pa- 



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287 



guridae and Parapaguridae. Only one her- 
mit crab has been reported to date from 
cold seeps of the Barbados accretionary 
prism, at depths of 1000-2000 m (K. Olu, 
pers. obs.), and another one from Monterey 
Bay seeps at 600-1000 m (J. P. Barry, pers. 
comm.). The Barbados specimen seems to 
belong to the genus Parapagurus (M. de 
Saint Laurent, pers. comm.). Although the 
Monterey Bay hermit crab has not been 
identified, it is possible that it is also a par- 
apagurid, which are common at these 
depths. 

Family Lithodidae 

The first report of the occurrence of Lith- 
odidae in hydrothermal vent or cold-seep 
habitats, is a "large crab," clearly a lith- 
odid, illustrated in a sketch of the benthic 
community associated with cold-seeps 
along the lower Oregon subduction zone at 
2037 m depth (Suess et al. 1985, Kulm et 
al. 1986). Subsequently, Carey et al. (1988) 
observed Lopholithodes foraminatus from a 
shallower part of this subduction zone. 
Whether the "large crab" and L. forami- 
natus are the same species is not stated. 
Tunnicliffe & Jensen (1987) proposed that 
the deeper water species from the lower 
zone could be the same Paralomis sp. they 
found at hydrothermal vents in the Juan de 
Fuca Ridge. 

In total, four lithodid crabs have been 
identified to the species level from cold- 
seep areas, and the status of four others re- 
mains uncertain. The Lithodidae are repre- 
sented at hydrothermal vents by two species 
and six occurrences not yet clearly assigned 
to species. Two of the described species 
{Paralomis arethusa and P. jamsteci) are 
until now known only from reducing envi- 
ronments. No lithodid species is definitely 
known from both vents and seeps. 

Genus Lithodes Latreille, 1806 
Lithodes manningi Macpherson, 1988 

One specimen was collected at 1236 m 
depth in cold seeps of the Barbados accre- 



tionary prism, and was identified by Mac- 
pherson (1994). The species is known from 
depths of 640-777 m in the Caribbean 
(Macpherson 1988). 

Two species of lithodid crabs found at 
seeps of the Monterey Bay, at 600-1000 m, 
have been assigned tentatively to the genus 
Lithodes (J. P. Barry, pers. comm.). These 
crabs are not considered as seep-endemics. 

Genus Lopholithodes Brandt, 1848 

Lopholithodes foraminatus (Stimpson, 

1859) 

As previously mentioned, Carey et al. 
(1988) observed dense aggregations of Lo- 
pholithodes foraminatus while exploring 
apparently extinct seepage areas of the up- 
per Oregon subduction zone, at a depth of 
ca. 250 m. There is no mention of this spe- 
cies at active seeps, but it could be the same 
species as the "large crab" observed at 
2037 m by Suess et al. (1985). However, 
this occurrence would be much deeper than 
the known depth range of L. foraminatus, 
which is known from British Columbia to 
southern California, at 0-547 m (Hart 
1982). 

Genus Neolithodes Milne Edwards & 

Bouvier, 1894 

Neolithodes diomedeae (Benedict, 1894) 

This species represents the first reported 
occurrence of Lithodidae in hydrothermal 
vents. Grassle (1986) described it as the 
most common crab found at the active hy- 
drothermal vents, at 2000 m, in the Guay- 
mas Basin, Gulf of California. It is known 
from southern California to South Georgia, 
at depths of 640-2450 m (Macpherson 
1988). 

Genus Paralithodes Brandt, 1848 

Based on submersible observations, Sa- 
galevich et al. (1992) reported Paralithodes 
sp., at 350-400 m, on the periphery of ac- 
tive hydrothermal vents on the summit of 
Piyp Volcano, in the Bering Sea, on dense 
populations of actinians. 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Genus Paralomis White, 1856 

This is the best represented of the Hth- 
odid genera in chemosynthetically-based 
communities. At least four species have 
been reported, including two exclusively 
from reducing habitats. Considering the nu- 
merous reports of ''Paralomis sp." found 
in the literature, more species are likely to 
be discovered in the future. 

Paralomis arethusa Macpherson, 1904 

Paralomis arethusa is one of two lith- 
odid crabs known from cold-seep commu- 
nities of the Barbados accretionary prism at 
1691 m depth. This species is known only 
from this location (Macpherson 1994). 

Paralomis cubensis Chace, 1939 

Sassen et al. (1993) produced a photo- 
graph of a "crab" (clearly a lithodid) crawl- 
ing on tubeworms and mussels at the Green 
Canyon methane seep, on the upper conti- 
nental slope of the Gulf of Mexico, at 620 
m. It was subsequently identified as Para- 
lomis cubensis. It is considered as a "va- 
grant" species of the Gulf of Mexico seeps 
by Carney (1994), and is not commonly in 
direct contact with the seep community. P. 
cubensis is known from east Florida to Bra- 
zil, at 329-730 m (Macpherson 1988). 

Paralomis jamsteci Takeda & Hashimoto, 
1990 

In hydrothermal areas of the Okinawa 
Trough, lithodid crabs have been observed 
on the Minami-Ensei Knoll, at 700 m, 
where a new species was described as Par- 
alomis jamsteci (Hashimoto et al. 1990, 
Takeda & Hashimoto 1990). This species is 
living among mytilid beds, near vent open- 
ings (Hashimoto et al. 1995). Two other yet 
unidentified species of Paralomis also oc- 
cur at the Minami-Ensei vents (Hashimoto 
et al. 1995). 



Paralomis multispina (Benedict, 1894) 

Horikoshi & Ishii (1985), Hashimoto et 
al. (1987, 1989), and Ohta (1990b) have de- 
scribed cold-seep communities of Sagami 
Bay in Japan, at depths of 900-1200 m, 
where large Paralomis multispina and the 
clams Calyptogena soyoae are the dominant 
species. P. multispina is known from Japan 
to Cahfornia, at 500-1665 m (Hart 1982). 

Paralomis verrilli (Benedict, 1894) 

Paralomis verrilli is believed to be the 
species found at hydrothermal vents of the 
Iheya Ridge, at 1400 m, in the Mid-Oki- 
nawa Trough (S. Ohta, pers. comm.; Ohta 
1990a; Kim & Ohta 1991; Hashimoto et al. 
1995). Although still to be confirmed, this 
occurrence would not be surprising as P. 
verrilli is usually found at depths of ca. 
1500-3500 m around Japan and from the 
Bering Sea to California (S. Ohta, pers. 
comm.). Crabs tentatively assigned to this 
species were also reported from the Sagami 
Bay cold-seeps, but in much lower numbers 
than the dominant P. multispina (J. Hashi- 
moto, pers. comm.). 

Additional occurrences o/ Paralomis spe- 
cies. — Several authors have reported the 
occurrence of lithodid crabs as belonging to 
the genus Paralomis. Most reports have 
been based on collected specimens. Two yet 
unidentified species of Paralomis occur at 
the Minami-Ensei vents (Mid-Okinawa 
Trough), living on mytilids or around bac- 
terial mats, together with the vent endemic 
P. jamsteci (Hashimoto et al. 1995). In the 
hydrothermal areas of the North-Fiji back- 
arc basin (2000 m) the "white-coloured 
lithodid crabs" observed by Jollivet et al. 
(1989), were later identified as belonging to 
the genus Paralomis, and have also been 
sampled at 1 800 m in venting areas of the 
Lau back-arc basin (Desbruyeres et al. 
1994). On the Juan de Fuca ridge in the 
northern Pacific, the lithodid crab observed 
and sampled at 1600 m on the Axial Sea- 
mount hydrothermal vent area by Tunni- 
cliffe & Jensen (1987), is a species of Par- 



VOLUME 109, NUMBER 2 



289 



alomis that co-occurs with the brachyuran 
majid crab Macroregonia macrochira. 
These authors suggested that this lithodid 
might be the same species found at the Or- 
egon cold-seeps by Suess et al. (1985). This 
Paralomis sp. and Paralomis verrilli (if 
confirmed) would then be the only lithodid 
species that occur in both vent and seep en- 
vironments. Galkin & Moskalev (1990b) 
also observed a crab closely related to Par- 
alomis on the Juan de Fuca ridge. 

Remarks on Lithodidae. — The Lithodi- 
dae do not seem to be physiologically de- 
pendent on the chemosynthetic production 
of vents or seeps. E. Escobar-Briones (pers. 
comm.) recently observed that although 
Neolithodes diomedeae was seen near bac- 
terial mats and hydrothermal vent structures 
in the Guaymas Basin, it appears to mainly 
feed on organic matter of photoautotrophic 
origin. The opportunistic behavior of this 
deep-sea scavenger is, however, believed to 
contribute to the export of hydrothermal 
material to the surrounding abyssal environ- 
ment. In other hydrothermal and cold-seep 
settings, direct evidence exists of lithodid 
crabs feeding on chemoautotrophic symbi- 
ont-containing fauna. Tunnicliffe & Jensen 
(1987) reported that pieces of vestimentif- 
eran tubes were found in the stomachs of 
Paralomis sp. at Axial Seamount vents. Ob- 
servations of P. multispina and P. jamsteci 
feeding upon live vesicomyid and mytilid 
bivalves are mentioned by Hashimoto et al. 
(1989) and Takeda & Hashimoto (1990) in 
Sagami Bay cold-seeps and Okinawa vents. 
Suess et al. (1985) described a large lith- 
odid in the Oregon seeps as actively feeding 
on vesicomyid clams. The Lithodes spp. 
from Monterey Bay seep communities are 
known to have attacked unsuccessfully ves- 
icomyid clams (J. P. Barry, pers. comm.). 
Another type of feeding has been proposed 
for P. multispina by Horikoshi & Ishii 
(1985), who observed this crab actively 
scooping up and "feeding" on the black re- 
duced mud within the clam beds of Sagami 
Bay cold-seeps. Most frequently, the Lith- 
odidae occur in greatest densities at vents 



and seeps. As there appears to be no other 
advantage in getting close to such a toxic 
milieu than the increased food-supply (par- 
ticularly near hydrothermal vents with high 
concentrations of hydrogen sulfide and 
heavy metals, low oxygenation and pH), 
they can be considered at least to partially 
feed on chemosynthetically-derived mate- 
rial. 

Cold-seep Lithodidae described to date 
range in depths from 250 to 2037 m. How- 
ever, one occurrence is known from cold- 
seep communities of the Nankai Trough, Ja- 
pan subduction zone, at 3800 m (M. Sibuet, 
pers. comm.). At hydrothermal vents, the 
depth range is 350-2000 m. The geograph- 
ical distribution of these observations is 
worldwide, with some noticeable excep- 
tions: no Lithodidae have ever been ob- 
served near hydrothermal vent areas of the 
East Pacific Rise (EPR), the Galapagos 
Rift, nor the Mid- Atlantic Ridge. The crab 
mentioned by Guinot & Macpherson (1987) 
from the EPR at 12°35'N, although proba- 
bly a lithodid, cannot be assigned to a hy- 
drothermal context. It was indeed observed 
on an off-axis seamount located 18 km 
from the ridge axis and even farther from 
the nearest known vent site in this area (P. 
Chevaldonne, pers. obs.). 

Superfamily Galatheoidea 

This superfamily includes four families, 
two of which are represented at deep-sea 
hydrothermal vents and cold seeps. 

Family Galatheidae 

Galatheid crabs have been found associ- 
ated with hydrothermal environments since 
the first submersible expeditions explored 
the Galapagos Rift vents (Corliss & Ballard 
1977). They have also been known to occur 
in cold-seep settings since the discovery of 
this type of community (Paull et al. 1984). 
Galatheids are familiar members of vent 
and seep communities but often do not at- 
tract as much attention as other spectacular 
organisms such as the Vestimentifera and 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



large bivalves. Also, because most gala- 
theid crabs appear similar, to the non-spe- 
cialists, observations on them are numerous 
but often vague. Only three of the 1 1 defi- 
nite species known so far have been re- 
ported from cold-seep communities, and 
one, Munidopsis crassa, from both hydro- 
thermal and seep environments. Reports of 
the latter, however, seem suspect as one is 
from the Mid-Atlantic Ridge, and another 
from the Peru margin. Many occurrences 
are mentioned only as "galatheid crabs," so 
much more species are likely to be de- 
scribed in the future. 

Genus Munida Leach, 1820 

Munida magniantennulata Baba & 

Turkay, 1992 

This species was originally described 
from three specimens collected at hydro- 
thermal vents of the Lau back-arc basin in 
the southwest Pacific, at a depth of 1750- 
2000 m (Baba & de Saint Laurent 1992, 
Baba & Turkay 1992). It was subsequently 
found in collections from a non-vent envi- 
ronment off Australia (Baba 1994). 

Munida microphthalma Milne Edwards, 
1880 

In the seepage areas of the Barbados ac- 
cretionary prism, at 1700-2000 m, two dif- 
ferent species of Galatheidae have been ob- 
served (K. Olu, pers. obs.). Only one has 
been collected and identified as M. micro- 
phthalma (M. de Saint Laurent, pers. 
comm.). Jollivet et al. (1990) earlier re- 
ported Munidopsis sp. from the same area. 
They either misidentified M. microphthal- 
ma from their towed camera pictures, or ob- 
served the second and bigger species, 
which might be a Munidopsis. Munida mi- 
crophthalma has been reported from the At- 
lantic, south of Iceland to Ascencion Island, 
in the Gulf of Mexico, the Carribbean, and 
the Bay of Biscaye, at depths of 194-2129 
m (Wenner 1982). 



Genus Munidopsis Whiteaves, 1874 

This is the most commonly encountered 
galatheid genus in reducing environments, 
including at least 10 species, of which six 
are known so far only from hydrothermal 
vents. Many authors have reported Muni- 
dopsis sp. or Munidopsis-like galatheid 
crabs, so again, the list of species is likely 
to expand. 

Munidopsis alvisca Williams, 1988 

This so far exclusively hydrothermal-as- 
sociated species was described by Williams 
(1988) from widely separated venting areas. 
Munidopsis alvisca occurs at the periphery 
of hydrothermal areas on the Explorer and 
Juan de Fuca ridges, at 1545-1800 m, in 
the northeastern Pacific, and at 2000 m in 
the Guaymas Basin, Gulf of California 
(Khodkina 1991). Van Dover et al. (1990) 
also reported a galatheid crab which might 
be M. alvisca, from vents of the Escanaba 
Trough on the Gorda Ridge. 

Munidopsis beringana Benedict, 1902 

A galatheid sampled from cold-seeps of 
the Middle America Trench at 3700-4000 
m west of Mexico has been assigned to 
Munidopsis beringana (E. Macpherson, 
pers. comm.). The galatheid crabs observed 
at shallower seep communities (2900 m) 
might belong to this same species (K. Olu, 
pers. obs.). There is still some doubt on the 
determination of this species, as the col- 
lected specimens present some variations 
from other M. beringana. This occurrence 
is also out of the known geographical and 
depth range of this species, usually found 
from the Bering Sea to Oregon, at 2800- 
3276 m (Ambler 1980). 

Munidopsis crassa Smith, 1885 

As previously mentioned, this species is 
believed to be present both in cold-seep and 
hydrothermal settings. Segonzac (1992) re- 
ported it from the Snake Pit hydrothermal 
area, at 3480 m, on the Mid- Atlantic Ridge. 



VOLUME 109, NUMBER 2 



291 



Olu et al. (1996) recently reported Muni- 
dopsis crassa from cold-seeps of the Peru 
margin, at 3000-3600 m, although uniden- 
tified galatheids were also observed at 5040 
m. A comparative examination of Atlantic 
and Peru specimens is needed to confirm 
these findings. M. crassa was previously 
only known from the Atlantic, at 2679- 
5315 m (Wenner 1982). 

Munidopsis diomedeae (Faxon, 1895) 

Munidopsis diomedeae was observed 
near hydrothermal vents of the Guaymas 
Basin, Gulf of California, at 2000 m depth 
(Khodkina 1991), along with M. alvisca. 
One of these two species might be the Mun- 
idopsis sp. mentioned by Lutz (1992) from 
cold seepages in the same area. M. diome- 
deae is common in the Gulf of California, 
and is known from California to Chile, at 
768-3790 m (Haig & Wicksten 1975). 

Munidopsis lauensis Baba & de Saint 
Laurent, 1992 

Munidopsis lauensis occurs with Munida 
magniantennulata at deep-sea vents of the 
Lau back-arc basin, in southwest Pacific, at 
1750 m. It is also found at 2000 m in hy- 
drothermal sites of the nearby North-Fiji 
back-arc basin (Baba & de Saint Laurent 
1992). To date, this species is exclusively 
known from hydrothermal areas. 

Munidopsis lentigo Williams & Van 
Dover, 1983 

This hydrothermal vent endemic is con- 
sidered a "high temperature" species (Wil- 
liams & Van Dover 1983) as it seems to be 
the galatheid that lives closest to high tem- 
perature fluid venting. It has been found 
only at vent sites of the East Pacific Rise at 
21°N, 2600 m depth. 

Munidopsis marianica Williams «& Baba, 
1989 

Munidopsis marianica is another appar- 
ently vent-endemic from the 3600-3700 m 



deep hydrothermal sites of the Mariana 
back-arc basin, western Pacific, where it 
lives in 10-25°C waters among dense car- 
pets of actinians (Williams & Baba 1989, 
Hessler & Lonsdale 1991). 

Munidopsis sonne Baba, 1995 

Munidopsis sonne is only known from 
two specimens collected at active vents of 
the North Fiji Basin, at 1992 m (Baba 
1995). 

Munidopsis starmer Baba & de Saint 
Laurent, 1992 

Munidopsis starmer also seems restricted 
to hydrothermal vents (Baba & de Saint 
Laurent 1992) and occurs at vent sites of 
the North-Fiji Basin, at 2750 m. In the same 
area, M. lauensis, M. sonne, and two spe- 
cies of Chirostylidae are also present. 

Munidopsis subsquamosa Henderson, 
1885 

Munidopsis subsquamosa is believed to 
be a complex of deep-sea cosmopolitan 
species (Van Dover 1986), which explains 
its considerable geographic range. Although 
this complex clearly needs a complete re- 
examination and taxonomic work at the 
molecular level, we will simply call it here 
M. subsquamosa. It was the first galatheid 
species observed in a chemosynthetically- 
based community, and is found at almost 
every vent location in the eastern Pacific, 
in a depth of ca. 2600 m (de Saint Laurent 
1984, Hessler et al. 1985, Van Dover et al. 
1985). At 21°N on the East Pacific Rise 
(EPR), it occurs with M. lentigo, but inhab- 
its colder waters. It is the only species ob- 
served at 13°N (EPR) and sites of the Ga- 
lapagos Rift, and has also been collected 
from the EPR at 10-12°N (Van Dover & 
Hessler 1990) and 9°50'N (Kaartvedt et al. 
1994). At the latter location, Lutz (1992) 
suggested that there might be two different 
species of Munidopsis. 

Additional occurrences of Munidopsis 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



species. — Many reports of ''Munidopsis 
sp." exist from a great variety of hydro- 
thermal and seepage areas. Hydrothermal 
occurrences include the EPR at 17-20°S 
(Van Dover & Hessler 1990); three loca- 
tions in the Mid-Okinawa back-arc basin, 
the Iheya Ridge at 1400 m (Ohta 1990a, 
Kim & Ohta 1991), the Izena Caldron at 
1400 m (Hashimoto et al. 1995), and the 
Minami-Ensei Knoll at 700 m (Hashimoto 
et al. 1990, 1995); Manus back-arc basin in 
the southwest Pacific at 2500 m (Galkin 
1992); TAG area on the Mid- Atlantic Ridge 
(Galkin & Moskalev 1990a); and Broken 
Spur area farther north on this same ridge 
(Murton et al. 1995). Reports from cold- 
seeps include the Nankai Trough in the Jap- 
anese Trench at 3800 m (Laubier et al. 
1986), cabonate seeps of Enshu-nada, Ja- 
pan, at 1000-1220 m (Ohta et al. 1995), 
and the Laurentian Fan at 3850 m (Mayer 
et al. 1988). 

In Monterey Bay, cold-seep communities 
occur at two different bathymetric levels, at 
3000-3600 m (Embley et al. 1990) and 
600-1000 m (J. R Barry, pers. comm.). At 
least two different species have been ob- 
served so far, possibly of two different gen- 
era. At deeper sites, a single species has 
been found and tentatively assigned to the 
genus Munidopsis by Embley et al. (1990), 
but could also be a species of Munida ac- 
cording to J. P. Barry (pers. comm.). At 
shallower seeps (600-1000 m), two differ- 
ent species of Galatheidae are known, one 
of which might be the same species ob- 
served deeper, and the other apparently is a 
seep endemic that might belong to the ge- 
nus Munidopsis (J. P. Barry, pers. comm.). 

In the Gulf of Mexico seep communities, 
MacDonald et al. (1989) reported a possibly 
undescribed species of Munidopsis from the 
Bush Hill site, at 540 m, on the Louisiana 
Slope. At different seepage depths on the 
slope (400-1000 m and 2200 m), Carney 
(1994) indicated two Munidopsis spp. (in- 
cluding one seep endemic), different from 
the "background" galatheids. One of these 
is probably the Bush Hill species. Muni- 



dopsis spp. are much more common than 
the chirostylid Eumunida picta that also oc- 
curs at the 400-1000 m sites, and they live 
in close association with mytilid bivalves 
and vestimentiferan tube worms. At 3270 m, 
in cold-seeps of the Florida Escarpment, 
Hecker (1985) has reported a large and nu- 
merous white Munidopsis sp. that has not 
been seen far outside the seep communities. 
Unconfirmed reports of Munidopsis-\\ke, 
galatheids include those from the hydro- 
thermal vents of the Piyp volcano, at 800 
m, in the Bering Sea (Sagalevich et al. 
1992). In the region of the Barbados accre- 
tionary prism, Munidopsis -like galatheids 
have also been observed associated with 
cold seeps on mud volcanoes, at 4710- 
4980 m (K. Olu, pers. obs.), and galatheids 
tentatively assigned to Munidopsis were re- 
ported by Jollivet et al. (1990), at shallower 
cold-seep sites (1000-2000 m), in the 
southern part of the prism. 



Family Chirostylidae 

Chirostylid crabs are represented in che- 
mosynthetic communities by two genera 
and three species. This family is frequently 
overlooked by non-specialist observers 
when reporting the occurrence of "gala- 
theids." 



Genus Eumunida Smith, 1883 
Eumunida picta Smith, 1883 

This species is considered as a rare va- 
grant in cold-seeps of the Louisiana slope, 
at 400-1000 m depth (Carney 1994). It has 
been reported from Massachusetts to Co- 
lombia, at 200-600 m (de Saint Laurent & 
Macpherson 1990). 



Genus Uroptychus Henderson, 1 



Baba & de Saint Laurent (1992) have de- 
scribed two new species only known from 
vents of the North-Fiji Basin. 



VOLUME 109, NUMBER 2 



293 



Uroptychus bicavus Baba & de Saint 
Laurent, 1992 

Uroptychus bicavus is known from a 
depth of 2750 m. 

Uroptychus thermalis Baba & de Saint 
Laurent, 1992 

Uroptychus thermalis occurs at depth of 
2000 m. 

Additional occurrences of Galatheo- 
idea. — Several authors have reported the 
occurrence of "galatheid crabs" or "squat 
lobsters" that cannot be assigned with cer- 
tainty to a family. These reports are from 
hydrothermal areas of the EPR at 21°30'S 
(Renard et al. 1985), 23°30'S and 26°S 
(Marchig & Gundlach 1987), the Gulf of 
Aden (Juniper et al. 1990) and the Edison 
Seamount in the southwestern Pacific (Her- 
zig et al. 1994), and from cold seeps of En- 
shu-nada, Japan, at 1000-1220 m (Ohta et 
al. 1995). 

Remarks on Galatheidae and Chirostyli- 
dae. — Like Lithodidae, Galatheidae and 
Chirostylidae are generally not considered 
to be dependent on the chemically reduced 
fluids emitted at vents and seeps. Their rep- 
resentatives are often restricted to the pe- 
riphery of the sites, where they are believed 
to be opportunistic species taking advantage 
of increased productivity. However, there is 
a significant number of species that seem to 
be endemic to hydrothermal vents or cold- 
seeps, and some of them even appear to be 
adapted to this harsh environment, not just 
the periphery. Van Dover (1986) observed 
that gravid females of the hydrothermal 
Munidopsis subsquamosa seemed to live 
under higher temperature conditions than 
males. Although this species is not restrict- 
ed to vent areas, this behaviour was inter- 
preted as a possible adaptation to improve 
reproductive efficiency. Galatheids also 
seem to adapt relatively well to hypoxic 
conditions, such as those encountered in re- 
ducing habitats. In some British Columbia 
fjords, for instance, dense aggregations of 
Munida quadrispina are consistently found 



at low oxygen levels, where most other in- 
vertebrates of the fjords cannot compete for 
food and space (Burd & Brinkhurst 1984). 
Stable isotope analyses led Van Dover & 
Fry (1989) and E. Escobar-Briones (pers. 
comm.) to characterize Munidopsis sub- 
squamosa and M. alvisca, of the Galapagos 
and Guaymas hydrothermal vents, as hav- 
ing a mixed diet partly including a sulfur- 
based source of carbon. Similar analyses on 
the Iheya Ridge Munidopsis sp. also show 
at least a partial nutritional dependence on 
chemosynthetic processes (Kim et al. 1989, 
Kim & Sakai 1991). At the Monterey Bay 
cold-seeps, an apparently seep endemic gal- 
atheid is found grazing on mats of the fil- 
amentous bacteria Beggiatoa, and is even 
covered with the bacterial filaments (J. R 
Barry, pers. comm.). 

Galatheids and chirostylids might also 
act as vectors for the export of organic mat- 
ter of hydrothermal origin in the deep sea. 
They might disperse more efficiently than 
lithodids, due to their swimming abilities, 
and can also easily detect new food sources. 
Two shark carcasses placed 100 m from a 
vent site at 13°N on the EPR (Fig. 1) at- 
tracted hundreds of Munidopsis subsqua- 
mosa within a day (D. Desbruyeres, pers. 
comm.). These characteristics might also 
favor dispersal of non-planktonic develop- 
mental stages of vent and seep species. Car- 
ney (1994) reported that Munidopsis sp. of 
the Gulf of Mexico seepage areas are often 
covered with juvenile mussels, and may be 
an important means of post-settlement dis- 
persal for seep endemic sessile taxa. Such 
phoretic processes might also occur in hy- 
drothermal polychaetes (Tunnicliffe & Jen- 
sen 1987, Zal et al. 1995). 

Galatheidae are found in virtually every 
hydrothermal or cold-seep environment, as 
well as in other types of reducing habitats 
such as whale carcasses (Bennett et al. 
1994, Wada et al. 1994) or decomposing 
wood (Wilhams & Baba 1989). They do not 
seem to be limited by depth, as their oc- 
currence is reported from as deep as 5040 
m, in the deepest cold-seep communities 



294 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




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VOLUME 109, NUMBER 2 



295 



(Olu et al. 1996). They are also associated 
with the shallowest occurrence of vestimen- 
tiferans known to date, at 82 m, in the cold- 
seep site of Kagoshima Bay, Japan (Hash- 
imoto et al. 1993). 

Acknowledgments 

We thank D. Desbruyeres, M. Sibuet, M. 
Segonzac and anonymous referees for help- 
ful comments on this manuscript, and A. 
Gebruk, S. Ohta, J. Hashimoto, J. P. Barry, 
and E. Escobar-Briones for communicating 
valuable information. R. Lemaitre, T. Com- 
tet, M. de Saint Laurent and D. Guinot pro- 
vided much appreciated help. 

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Bulletin of the Biological Society of Washing- 
ton 6:475-484. 

Takeda, M., & J. Hashimoto. 1990. A new species of 
the genus Paralomis (Crustacea, Decapoda, 
Lithodidae) from the Minami-Ensei Knoll in the 
mid-Okinawa trough. — Bulletin of the National 
Science Museum, Tokyo, Series A 16:79-88. 

Tunnicliffe, V., & R. G. Jensen. 1987. Distribution 
and behaviour of the spider crab Macroregonia 
macrochira Sakai (Brachyura) around the hy- 
drothermal vents of the northeast Pacific. — Ca- 
nadian Journal of Zoology 65:2443-2449. 

Van Dover, C. L. 1986. A comparison of stable iso- 
tope ratios ('*0/'*0 and '^C/'^C) between two 
species of hydrothermal vent decapods {Alvi- 
nocaris lusca and Munidopsis subsquamosa). — 
Marine Ecology Progress Series 31:295-299. 

, J. R. Factor, A. B. Williams, & C. J. Berg Jr. 

1985. Reproductive patterns of decapod crus- 
taceans from hydrothermal vents. — Bulletin of 
the Biological Society of Washington 6:223- 
227. 



— , & B. Fry. 1989. Stable isotopic compositions 
of hydrothermal vent organisms. — Marine Bi- 
ology 102:257-263. 

-, J. E Grassle, & M. Boudrias. 1990. Hydro- 
thermal vent fauna of Escanaba Trough (Gorda 
Ridge). Pp. 285-287 in G. R. McMurray, ed., 
Gorda Ridge: a seafloor spreading center in the 
United States Exclusive Economic Zone. 
Springer- Verlag, New York. 

-, & R. R. Hesslen 1990. Spatial variation in 



faunal composition of hydrothermal vent com- 
munities on the East Pacific Rise and Galapagos 
spreading center. Pp. 253-264 in G. R. Mc- 
Murray, ed., Gorda Ridge: a seafloor spreading 
center in the United States Exclusive Economic 
Zone. Springer- Verlag, New York. 

Wada, H., T. Naganuma, K. Fujioka, H. Kitazato, K. 
Kawamura, & Y. Akazawa. 1994. The discov- 
ery of the Torishima Whale Bone Animal Com- 
munity (TOWBAC) and its meaning: the results 
of revisit dives by the "Shinkai 6500." — JAM- 
STEC Journal of Deep Sea Research 10:37-47 
(in Japanese with English summary). 

Wenner, E. L. 1982. Notes on the distribution and 
biology of Galatheidae and Chirostylidae (De- 
capoda: Anomura) from the Middle Atlantic 
Bight. — Journal of Crustacean Biology 2:360- 
377. 

White, A. 1856. Some remarks on Crustacea of the 
genus Lithodes with a brief description of a spe- 
cies apparently hitherto unrecorded. — Proceed- 
ings of the Zoological Society of London 1856: 
132-135. 

Whiteaves, J. E 1874. On recent deep-sea dredging 
operations in the Gulf of St. Lawrence. — Amer- 
ican Journal of Science, Series 3 7:210-219. 

Williams, A. B. 1988. New marine decapod crusta- 
ceans from waters influenced by hydrothermal 
discharge, brine, and hydrocarbon seepage. — 
Fishery Bulletin 86:263-287. 

, & K. Baba. 1989. New squat lobsters (Gal- 
atheidae) from the Pacific Ocean: Mariana 
back-arc basin. East Pacific Rise, and Cascadia 
Basin.— Fishery Bulletin 87:899-910. 

, & C. L. Van Dover. 1983. A new species of 



Munidopsis from submarine thermal vents of 
the East Pacific Rise at 21°N (Anomura: Gal- 
atheidae). — Proceedings of the Biological So- 
ciety of Washington 96:481-488. 
Zal, E, D. JoUivet, P. Chevaldonne, & D. Desbruyeres. 
1995. Reproductive biology and population 
structure of the deep-sea hydrothermal vent 
worm Paralvinella grasslei (Polychaeta: Alvi- 
nellidae) at 13°N on the East Pacific Rise. — 
Marine Biology 122:637-648. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):299-305. 1996. 

A new species of Aniculus Dana (Decapoda: Anomura: Diogenidae) 

from Hawaii 

Patsy A. McLaughlin and John P. Hoover 

(PMcL) Shannon Point Marine Center, Western Washington University, 

1900 Shannon Point Road, Anacortes, Washington 98221-4042, U.SA. 

(JPH) Hawaii Medical Library, 1221 Punchbowl Street, 

Honolulu, Hawaii 96813, U.S.A. 

Abstract. — A new and spectacular, but rarely seen, species of the hermit crab 
genus Aniculus Dana (Aniculus hopperae) is described and illustrated. It is 
compared and contrasted with Aniculus maximus Edmondson, the only other 
species of the genus known from the Hawaiian Islands and with A. retipes 
Lewinsohn, a species to which it is most morphologically related. 



While diving at Pupukea on the north 
shore of the Hawaiian island of Oahu dur- 
ing June of 1995, the junior author collect- 
ed an unfamiliar hermit crab inhabiting a 
drupe shell, Drupa rubusidaeus Roding, 
1798 (aperture length about 2 cm). At a 
depth of about 8 m, the shell lay on boulder 
and rubble bottom adjacent to the low lime- 
stone cliffs of the shoreline containing a 
system of deep undercuts and small caves. 

A brief literature search by the junior au- 
thor revealed no named species to which it 
could be referred. However, a photograph 
of a similar animal from a popular book on 
hermit crabs (Giwojna 1978: 57) bore the 
following caption: "This species of Dar- 
danus with red legs and claws and orange 
eye-stalks and antennae, hails from Ha- 
waii." 

The photographer, Scott Johnson, con- 
tacted by telephone, was able to locate an 
additional photograph of the species in 
question, taken at Makua, Oahu on a ledge 
at a depth of 8-10 m, a site on the western 
shore similar to that at Pupukea where the 
holotype was collected. 

The Pupukea specimen was kept alive in 
a plastic container full of seawater that was 
changed every other day until it could be 
photographed in natural surroundings, then 
preserved in 70% alcohol and sent to the 



senior author. It has now been deposited in 
the collection of the National Museum of 
Natural History, Smithsonian Institution 
(USNM). A second specimen, subsequently 
collected, has been deposited in the collec- 
tion of the Bemice P. Bishop Museum 
(BPBM). Shield length (SL), measured 
from the midpoint of the anterior carapace 
margin to the midpoint of the posterior mar- 
gin of the shield provides an indication of 
size. 

Aniculus hopperae, new species 
Figs. 1-3 

Dardanus species: Giwojna, 1978: 57, un- 
numbered figure. 

Holotype. — S (SL = 6.32 mm), Pupu- 
kea, Oahu, Hawaii, 8 m, 18 Jun 1995, coll. 
J. Hoover; USNM 275921. 

Paratype. — S (SL = 2.96 mm), Pupu- 
kea, Oahu, 10 m, 7 Oct 1995, coll. R. Hol- 
com; BPBM SI 1284. 

Description. — Shield (Figs. lA, 2A) lon- 
ger than broad, with few shallow depres- 
sions, but not divided into distinct lobes. 
Mesogastric region delimited posteriorly 
only by pair of furrows forming weak V; 
with deep transverse post-rostral furrow. 
Rostrum triangular, acute, approximately 
reaching level of well developed, spinose 



300 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




i A-D,F 



Fig. L Aniculus hopperae, new species. Holotype, USNM 275921: A, shield and cephalic appendages; B, 
carpus and chela of left cheliped (outer face); C, coxae of chelipeds (ventral view); D, third left pereopod (lateral 
view); E, anterior lobe of sternite of third pereopods; F, tergite of sixth abdominal somite and telson (dorsal 
view). Scales equal 3.0 mm (A-D, F) and 1.0 mm (E). 



VOLUME 109, NUMBER 2 



301 




B 


-.. ^x -^ -^'' ^" ' ■ ^» V. 






^ 






^ 




^T 


;J 









Fig. 2. Aniculus hopperae, new species. Holotype, USNM 275921: A, whole animal (dorsal view); B, left 
second and third pereopods (lateral view). Scales equal 4.0 mm (A) and 2.0 mm (B). 



302 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



lateral projections. Ocular peduncles un- 
equal in length, left longest, slightly longer 
than shield, bulbous basally, appreciably 
thinner medially; comeae not dilated. Oc- 
ular acicles very well developed, triangular, 
with simple terminal spine. 

Antennular peduncles reaching to distal 
third of ocular peduncles. Ultimate and pen- 
ultimate segments with scattered long, stiff 
setae. Basal segment with scattered stiff se- 
tae, and minute spinule on laterodistal mar- 
gin. 

Antennal peduncles shorter than anten- 
nular peduncles, reaching little beyond 
proximal half of ocular peduncles. Fifth, 
fourth and third segments unarmed, but 
with scattered long stiff setae, most abun- 
dant at ventrodistal margin of third. Second 
segment with dorsolateral distal angle pro- 
duced, with small terminal spine practically 
obscured by long stiff setae; dorsomesial 
distal angle unarmed, mesial margin with 
long stiff setae. First segment with 3 or 4 
spinules on ventrolateral margin distally. 
Antennal acicle subtriangular, elongate, 
reaching well beyond proximal margin of 
ultimate peduncular segment, with terminal 
spine obscured by tuft of long setae and ad- 
ditional tufts of similar setae mesially and 
laterally. Antennal flagellum long, over- 
reaching outstretched chelipeds and ambu- 
latory legs; each article with 1 or 2 minute 
setae. 

Chelipeds (Figs. IB, C, 2A, 3A, B) and 
ambulatory legs short. Chelipeds equal, 
similar; overreaching distal margin of cor- 
neae by half length of palm; tips of dactyls 
and fixed fingers reach slightly beyond 
proximal margins of dactyls of second per- 
eopods and mid-length of dactyls of third. 
Dactyls slightly longer than palm; cutting 
edges of both dactyl and fixed finger with 
few large calcareous teeth, terminating in 
very prominent corneous hoof-like claws; 
dorsal and mesial surfaces of dactyls each 
with transverse rows of striae armed with 
small corneous spines and stiff setae, con- 
tinuing onto ventral surfaces as rows of stri- 
ae set with short stiff setae. Palms with 



short, transverse rows of 1 to 4 corneous- 
tipped tubercles on dorsal surface in mesial 
fourth, remainder of palm and fixed finger 
with transverse striae almost completely 
circumscribing dorsal, lateral and mesial 
surfaces and set with rows of quite short 
setae ventrally and laterally, becoming lon- 
ger and supplemented with corneous spi- 
nules on dorsal surface of fixed finger and 
medianly on dorsal surface of palm. Carpi 
approximately equaling length of palms; 
striae of dorsal surfaces not disposed in reg- 
ular transverse lines but rather in network 
of curves inset with short stiff and few lon- 
ger stiff setae, dorsomesial distal angles 
each with 1 prominent corneous-tipped 
spine and 3 corneous spinules on dorsodis- 
tal margin; mesial, lateral and ventral sur- 
faces with transverse, sometimes short or 
interrupted, striae inset with short stiff se- 
tae. Meri subtriangular; dorsal margins with 
transverse rows of striae inset with short se- 
tae, extending onto lateral and ventral sur- 
faces, dorsally accompanied by tufts of long 
setae; ventroraesial margins each with 2 or 
3 very small spines distally, ventral surfaces 
at mesial angles each with cluster of cor- 
neous-tipped spinules. Ischia each with 2 
transverse striae inset with short setae on 
ventral surface. Coxae (Fig. IC) each with 
very short stria mesially. 

Second and third pereopods (Figs. ID, 
2B) similar from left to right, third pair 
slightly shorter than second; dactyls with 
tufts of long stiff setae on all surfaces, pro- 
podi, carpi, meri and ischia with tufts of 
long setae predominately dorsally and ven- 
trally. Dactyls 0.75 to 0.80 length of pro- 
podi; dorsal surfaces of both second and 
third, and mesial and lateral faces of third 
each with irregular rows of corneous spi- 
nules; ventral margins each with 7 or 8 cor- 
neous spines. Propodi slightly longer than 
carpi and approximately equal to length of 
meri; 1 continuous transverse stria inset 
with short setae circumscribing dorsal, lat- 
eral and ventral surfaces distally, followed 
posteriorly by series of striae interrupted 
medially on lateral surfaces; mesial faces 



VOLUME 109, NUMBER 2 



303 




Fig. 3. Aniculus hopperae, new species. Holotype, USNM 275921: A, chela and carpus of left cheliped 
(dorsal view); B, chela and carpus of right cheliped (dorsal view). Scales equal 2.0 mm. 



with scattered tufts of setae; ventrodistal 
margins each with 1 or 2 corneous spinules. 
Carpi with few transverse striae incom- 
pletely circumscribing dorsal, lateral and 
mesial surfaces; dorsodistal angles each 
with corneous-tipped spine. Meri each with 



transverse striae circumscribing dorsal, lat- 
eral and ventral surfaces, interrupted me- 
dianly on lateral faces; striae of second pair 
much less distinct than third; second each 
with small spine at ventrolateral distal an- 
gle. Anterior lobe of stemite of third per- 



304 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



eopods (Fig. IE) subrectangular, with long 
marginal setae. 

Tergites of abdominal somites 1-5 mem- 
braneous but clearly delineated and orna- 
mented with long greenish- or whitish- 
tipped red marginal setae; tergite of sixth 
(Fig. IF) with dorsal longitudinal median 
furrow of anterior lobe very weakly 
marked, continued on posterior lobe as 
ovate depression and longitudinal series of 
deep pits posteriorly. Telson (Fig. IF) with 
transverse suture not much more distinct 
that supplemental lateral notches; posterior 
lobes asymmetrical, subtriangular, left 
slightly to appreciably longer than right; 
with narrow median cleft; terminal margins 
with long setae, longest at external angles. 

Color. — Long setae overall red proxi- 
mally, greenish-white distally, short setae of 
striae greenish or yellowish-white; spines 
white with black corneous tips; claws and 
corneous spines black. Shield mottled red 
and white with purple tint medianly and an- 
terior to post-rostral furrow. Ocular pedun- 
cles yellow; ocular acicles mottled red and 
white, with white terminal spinule. Anten- 
nular peduncles and flagella light yellow. 
Proximal segments of antennal peduncle 
and acicle mottled red and white; ultimate 
segment and flagella light purple. Chelae of 
chelipeds brilliant red with scattered white 
specks and splotches. Carpi brownish red 
with specks and splotches of white. Meri 
with lighter patches of brownish-red or 
mottled red and white on white back- 
ground. Dactyls of ambulatory legs red 
with white patches. Propodi each with ir- 
regular longitudinal white band dorsally 
and ventrally; mesial faces mottled red and 
white; lateral faces red with white striae and 
median longitudinal band of purple. Carpi 
each with longitudinal white stripe on ven- 
tral surface; mesial faces mottled red and 
white; dorsal and lateral surfaces red, striae 
white, striae of lateral faces separated by 
longitudinal band of purple. Meri mottled 
red and white; longitudinal purple band of 
carpi continued on lateral faces dorsolater- 
ally. 



Distribution. — At present known only 
from two locations on Oahu, Hawaii; 8-10 
m. 

Etymology. — This species is named in 
honor of Dr. Carol N. Hopper, Director of 
Education at Honolulu's Waikiki Aquarium, 
an enthusiastic advocate of Hawaii's marine 
invertebrates, and an inspiration to the 
many young aspiring marine biologists who 
attend her classes. 

Affinities. — Aniculus hopperae is imme- 
diately distinguished from the other resident 
Hawaiian species of the genus, A. maximus 
Edmondson, 1952, by its color, the sculp- 
turing of the shield, the arrangement of stri- 
ae on the ambulatory legs, and the armature 
of the tergite of the sixth abdominal somite. 
In A. maximus, the general color is intense 
orange or golden yellow with the striae set 
off by red-violet, the setae are yellow and 
the bristles red-tipped; the shield is marked 
by a series of furrows that clearly delineate 
the rhomboid mesogastric region common 
to most species of Aniculus; the transverse 
striae on the lateral faces of the propodi of 
the ambulatory legs are continuous; and the 
anterior lobe of the tergite of the sixth ab- 
dominal somite is armed with a cluster of 
spinules on either side of the midline. In 
contrast, the general color of A. hopperae 
is brilliant red, the striae often are white and 
the long setae are red, tipped with greenish 
white; the shield lacks any clear delineation 
of the mesogastric region; the striae of the 
lateral faces of the propodi of the ambula- 
tory legs are interrupted by a broad smooth 
longitudinal band of purple; and the tergite 
of the sixth abdominal somite has an un- 
armed anterior lobe. 

Forest (1984:59) remarked that A. retipes 
Lewinsohn, 1982 "... occupe une position 
incontestablement isolee par rapport aux 
autres especes du genre." Those characters 
which set A. retipes apart included the lack 
of a delimited mesogastric region and the 
interruption of the striae on the lateral faces 
of the propodi of the ambulatory legs by a 
smooth longitudinal band. Aniculus hopper- 
ae shares both of these characters with A. 



VOLUME 109, NUMBER 2 



305 



retipes. Additionally the tergite of the sixth 
abdominal somite has an unarmed anterior 
lobe in both species; however, that is a 
character common to all species except A. 
maximus. Aniculus retipes and A. hopperae 
are readily distinguished by their color pat- 
terns, as well as the different patterns of the 
striae on the carpi of the chelipeds. The oc- 
ular peduncles of A. hopperae are uniform- 
ly yellow, where as those of A. retipes have 
longitudinal red stripes. Aniculus retipes, 
particularly specimens from Phuket, Thai- 
land, have appreciably longer antennular 
peduncles (Forest 1984:52, fig. 51) than 
seen in A. hopperae. 

Remarks. — Like A. hopperae, the major- 
ity of the seven other described species of 
Aniculus are known from relatively few 
specimens. Only A. aniculus (Fabricius, 
1787) and A. ursus (Olivier, 1811) can be 
considered common. However, most spe- 
cies have broad, albeit sporatic, distribu- 
tions. For example, the other Hawaiian spe- 
cies, A. maximus, has also been reported 
from the Marquesas and Seychelles Islands. 
Similarly, the closely related A. retipes is 
known from the Red Sea to Zanzibar in the 
western Indian Ocean, to the west coast of 
Malaysia in the east, and in the western Pa- 
cific from the South China Sea and Banda 
Sea of Indonesia to Samoa (Forest 1984). 

Twenty photographs of the living holo- 
type were distributed to members of the Ha- 
waiian Malacological Society, dive charter 
operators, Waikiki Aquarium staff and other 
likely acquaintances in the hope that further 
specimens and ecological data would come 
to light. Subsequently, John L. Earle, a 
long-time Oahu shell collector, informed 
the junior author that the crab was familiar 
to him, although not encountered very of- 
ten. He could not cite an exact location or 
depth but said that he usually collected in 



the rubble at the bottom of a drop-off near 
Makua, Oahu at a depth of about 23 m. The 
drop-off contains numerous undercuts and 
caves. In the absence of additional infor- 
mation, the authors postulate this species to 
be an uncommon inhabitant of exposed 
rocky Hawaiian shores at a depth of 8 to 
23 m or more, in the vicinity of caves and 
ledges (where it may shelter during the 
enormous winter surf)- It is almost certainly 
not an intertidal species; this biotope has 
been well sampled on Oahu for many years. 
Like many hermit crabs, it is probably most 
active at night. 

Acknowledgments 

The authors gratefully acknowledge the 
collection of the second specimen by Ron 
Holcom and the ecological information pro- 
vided by John Earle. This is a scientific 
contribution from the Shannon Point Ma- 
rine Center, Western Washington Universi- 
ty- 
Literature Cited 

Edmondson, C. H. 1952. Additional central Pacific 
crustaceans. — Occasional Papers of Bernice P. 
Bishop Museum, 21(6):67-86. 

Fabricius, J. C. 1787. Mantissa insectorum sistens 
eorum species nuper detectas adiectis character- 
ibus genericis, differentiis specificis, emenda- 
tionibus, observationibus. I:i-xx, 1-348, Haf- 
nia. 

Forest, J. 1984. Revision du genre Aniculus. — Crus- 
taceana. Supplement 8:1-91. 

Giwojna, P. 1978. Marine hermit crabs. T.F.H. Publi- 
cations, Neptune City, New Jersey, 128 pp. 

Lewinsohn, Ch. 1982. Aniculus retipes n. sp. ein 
neuer Einsiedlerkrebs aus dem Roten Meer (De- 
capoda, Anomura). — Crustaceana, 42(l):76-82. 

Olivier, A. G. 1811. Pagure. Pagurus. Encyclopedic 
methodique. — Histoire naturelle, Insectes, 8: 
631-647. 

Roding, P. F. 1798. Museum Boltenianum sive Catal- 
ogus Cimeliorum, pars secunda Conchylia. 
Hamburg, Trappii. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2): 306-3 10. 1996. 

Description of a new porcellanid, Petrolisthes gertrudae from the 
southeastern Caribbean Sea (Crustacea: Decapoda: Porcellanidae) 

Bernd Werding 

Institut fiir Allgemeine und Spezielle Zoologie der Justus-Liebig-Universitat, 
Stephanstr. 24 D-35390 Giessen, Germany 

Abstract. — A new species of the genus Petrolisthes Stimpson, 1858, P. ger- 
trudae, from the West Indies is described. The new species belongs to a group 
of Petrolisthes species characterized by the complete lack of spines on the 
carapace and pereopods 1-5. The new species can be easily distinguished from 
other species of that group by the convex shape of the carpal lobe of the 
chelipeds, and by the pubescence which covers the outer margins of the chelae. 



The porcellanid fauna of the Caribbean 
Sea can be considered fairly well known, 
and has been documented in a series of 
publications treating species from the 
southern Caribbean (Gore 1970, 1974, 
1982; Gore & Abele 1973, 1976; Werding 
1977, 1978a, 1978b, 1982, 1986). Werding 
(1986) considered the species inventory of 
the western Atlantic as almost complete. A 
total of 43 species is now known from the 
western Atlantic, including three endemics 
from the coast of Brazil. 

The study of collections in the National 
Museum of Natural History, Smithsonian 
Institution, revealed the presence of an un- 
described species from the southeastern Ca- 
ribbean. The specimens were collected by 
P. Hummelinck in 1931, at Bonaire (Dutch 
West Indies), and by W L. Schmitt in 1956, 
at Guadeloupe (French West Indies). 

All type specimens are deposited in the 
National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
(USNM). The measurements given refer to 
carapace length (CL) and carapace width 
(CW) respectively. 

Family Porcellanidae 

Petrolisthes gertrudae, new species 

Figs. 1, 2 

Material. — Holotype: ovigerous female, 
Guadeloupe, West Indies, Pointe a Pitre, 



Rat Islands, exposed reef, submerged 
weedy rock, Smithsonian Bredin Caribbean 
Expedition, 30 Mar 1956, leg. W L. 
Schmitt USNM 275965. Paratypes: 1 ovi- 
gerous female, same data as holotype 
USNM 275966; 1 ovigerous female, Bo- 
naire, West Indies, Kralandijk, in colonies 
of Zoanthus sociatus, 31 Oct 1930, leg. P. 
Hummelinck USNM 275967. 

Measurements. — Holotype: CL 3.38 mm, 
CW 3.51 mm. Paratype from Guadeloupe: 
CL 4.29 mm, CW 4.35 mm; paratype from 
Bonaire: CL 3.45 mm, CW 3.64 mm. 

Range of length/width of carpus of che- 
liped: 1.35-1.44; range of egg diameter: 
0.65-0.72 mm. 

Description. — Carapace subquadrate and 
about as long as broad; lateral margins 
nearly parallel behind epibranchial angle. 
Surface almost smooth, without setae, cov- 
ered anteriorly with fine granules and be- 
hind epibranchial angle with fine plicae; 
grooves marking regions distinct. Frontal 
region strongly deflexed, produced, slightly 
trilobate, median lobe broad, triangular with 
shallow median groove. Flanks between 
side walls and legs with fringe of plumose 
setae. Orbits shallow, outer orbital angle 
slightly produced. No epibranchial spine. 

Inner part of ocular peduncle expanded 
to a longitudinal crest. 

First movable segment of antennae an- 



VOLUME 109, NUMBER 2 



307 




E ^. 

CO ^ 



O 



M 

£ 



308 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 2. Petrolisthes gertrudae, new species. Female paratype from Guadeloupe, West Indies USNM 275966. 
a, left cheliped ventral view; b, right third maxilliped, external view; c, propodus and dactylus of first right 
walking leg, ventral view; d, basal segment of right antennula, ventral view. 



teriorly extended to form low, granular 
crest; second irregularly granulate; third 
nearly smooth. Flagellum thick, setose. 
Outer surface of ischium of third maxilliped 
slightly rugose. 

Chelipeds subequal; merus and carpus 
without setae, dorsal surface with large flat- 
tened granules. Merus with broad granular 
lobe on anterior margin. Carpus less than Vi 
as long as wide, anterior margin extended, 
forming rounded lobe, strongly convex at 
proximal %; proximal portion sometimes 
armed with fine, irregular denticulation 
along edge; with shallow longitudinal 



groove marking broad rounded crest on 
dorsal surface near the posterior margin, 
crest composed of large transversally ori- 
ented granules; outer distal angle rounded. 
Surface of manus with similar granulation, 
upper side with large longitudinal elevation 
extending to gape of fingers; outer border 
with longitudinal depression, forming 
rounded crest at outer margin; outer margin 
distally straight or slightly curved outward, 
pollex bent upward; with thick tuft of long 
feathered setae covering outer margin leav- 
ing free only proximal portion and distal 
half of pollex. Fingers curved at tips, cut- 



A 



VOLUME 109, NUMBER 2 



309 



ting edges without prominent teeth, fingers 
meeting for entire length or slightly gaping, 
gape lacking pubescence. 

Walking legs with surface granulation 
similar to that of chelipeds; unarmed, with 
scattered plumose setae. Propodus with 3 
movable spines distally, and 2 or 3 addi- 
tional ones on inner margin variing incon- 
sistently among legs. Dactylus with 4 mov- 
able spines on inner margin. 

Abdomen with dense fringe of plumose 
setae on entire outer margin; telson con- 
sisting of 7 plates. 

Variations. — The paratype specimen 
from Bonaire exhibits a more accentuated 
granulation on the surface of the carapace, 
and specially at the extremities, than the 
two specimens from Guadeloupe. The an- 
terior lobe of the carpus of the chelipeds is 
somewhat depressed by a longitudinal 
groove. 

Habitat. — The specimen collected by P. 
Hummelinck was found in a colony of the 
zoanthid Zoanthus sociatus Ellis, a species 
which settles on the wave surf zone of the 
rocky intertidal. The specimens by W. L. 
Schmitt were found on an exposed reef. 
The species seems to be confined to the up- 
per littoral. 

Etymology. — The species is named in 
honor of my wife, Mrs. Gertrud Werding, 
for her help and understanding during all 
the years that I have been working on crus- 
taceans. 

Discussion 

Petrolisthes gertrudae, new species, be- 
longs to a group of species which is mainly 
confined to the eastern Pacific, where it is 
represented by 17 tropical and temperate 
species (Haig 1960). The group is charac- 
terized by the complete lack of spines on 
the carapace and pereopods, including the 
chelipeds. The carpus of the chelipeds is 
unarmed, with the exception of minute spi- 
nules in some species. The group was rep- 
resented until now in the western Atlantic 
by P. quadratus Benedict, P. tridentatus 



Stimpson, and P. tonsorius Haig. The new 
species is clearly distinguished from all oth- 
er species of the group by the convex lobe 
of the anterior margin of the carpus of the 
chelipeds which, in some cases, is slightly 
dentate, and the dense pubescence on the 
outer border of the manus absent in other 
representatives of the group. 

Acknowledgments 

I am very grateful to Dr. Rafael Lemaitre 
for his help and orientation during my visit 
to the National Museum of Natural History, 
Washington, D.C., and to him and Dr. Roy 
K. Kropp for the revision of the manuscript. 
To the Smithsonian Institution's Office of 
Fellowships and Grants, I acknowledge 
support in the form of a short- term visitor 
grant. I am grateful to Mrs. Helga Schmitt 
for kindly inking the drawing of the type 
specimen. 

Literature Cited 

Gore, R. H. 1970. Pachycheles cristobalensis sp. nov., 
with notes on the porcellanid crabs of the south- 
western Caribbean. — Bulletin of Marine Sci- 
ence 20(4):957-970. 

. 1974. Biological results of the University of 

Miami Deep-Sea Expeditions, 102: on a small 
collection of porcellanid crabs from the Carib- 
bean Sea (Crustacea, Decapoda, Anomura). — 
Bulletin of Marine Science 24(3):700-721. 

. 1982. Porcellanid crabs from the coasts of 

Mexico and Central America (Crustacea: De- 
capoda: Anomura). — Smithsonian Contribu- 
tions to Zoology 363:1-34. 

, & L. G. Abele. 1973. Three new species of 

porcellanid crabs (Crustacea, Anomura) from 
the Bay of Panama and adjacent Caribbean wa- 
ters. — Bulletin of Marine Science 23(3):559- 
573. 

. 1976. Shallow water porcelain crabs from the 



Pacific coast of Panama and adjacent Caribbean 
waters (Crustacea: Anomura: Porcellanidae). — 
Smithsonian Contributions to Zoology 237:1- 
30. 

Haig, J. 1960. The Porcellanidae (Crustacea Anomu- 
ra) of the eastern Pacific. — Allan Hancock Pa- 
cific Expeditions 24:1-440, pi. 1-41. 

Stimpson, W. 1858. Prodromus descriptionis animal- 
ium evertebratoiTim, quae in expeditione ad 
Oceanum Pacificum septentrionalem, a repub- 
lica Federata Missa, Cadwaladaro Ringgold et 



310 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Johanne Rodgers Ducibus, observavit et des- 
cripsit, pars VII, Crustacea Anomura I: Teleo- 
somi. — Proceedings of the Academy of Natural 
Sciences of Philadelphia (1858) 10:225-252. 

Werding, B. 1977. Los porcelanidos (Crustacea, An- 
omura, Porcellanidae) de la region de Santa 
Marta, Colombia. — Anales del Institute de In- 
vestigaciones Marinas de Punta de Betin 9:173- 
214. 

. 1978a. Ein Porzellanide, Petrolisthes mag- 

dalenensis n.sp. von der karibischen Kiiste Ko- 
lumbiens (Crustacea, Decapoda, Anomura). — 
Senckenbergiana biologica. 59:307-310. 

. 1978b. Los porcelanidos (Crustacea: Ano- 
mura: Porcellanidae) de la region de Acandi 



(Golfo de Uraba) con algiinos encuentros nue- 
vos de la region de Santa Marta (Colombia). — 
Anales del Institute de Investigaciones Marinas 
de Punta de Betin 10:213-221. 

— . 1982. Porcellanid crabs of the Islas del Ro- 
sario, Caribbean coast of Colombia, with a de- 
scription of Petrolisthes rosariensis new species 
(Crustacea: Anomura). — Bulletin of Marine 
Science 32(2):439-447. 

— . 1986. Die Gattung Neopisosoma Haig, 1960 
im tropischen Westatlantik mit der Beschrei- 
bung von Neopisosoma neglectum spec. nov. 
und Neopisosoma orientale spec. nov. (Crusta- 
cea: Anomura: Porcellanidea). — Zoologische 
Mededelingen, Leiden 16(11):159-179. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):31 1-317. 1996. 

Nannotheres moorei, a new genus and species of minute pinnotherid 

crab from Belize, Caribbean Sea 

(Crustacea: Decapoda: Pinnotheridae) 

Raymond B. Manning and Dairy 1 L. Felder 

(RBM) Department of Invertebrate Zoology, National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 20560, U.S.A.; 

(DLF) Department of Biology, University of Southwestern Louisiana, 

Lafayette, Louisiana 70504, U.S.A. 

Abstract. — Nannotheres moorei is described from a small pteriid bivalve 
mollusk taken in the Caribbean Sea off Belize. This minute, ovigerous pin- 
notherid crab can be distinguished by its size and the 2-segmented palp on its 
third maxilliped. With a carapace width of about 1 .5 mm in a sexually mature 
female, it may be the smallest known species of crab. 



The minute crab described below was 
found on the edge of the shell of its host 
by Donald R. Moore, University of Miami, 
who gave it to us for study. The unique 
holotype is in the collections of the Nation- 
al Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM). 

Abbreviations used below include: cb, 
carapace breadth; m, meters; mm, milli- 
meters; Mxp3, third maxilliped; P1-P5, 
pereopods (PI is the cheliped, P2-P5 the 
walking legs); Pip, pleopod. 

Nannotheres, new genus 

Diagnosis. — Adult female: size very 
small, carapace length less than 2 mm. Car- 
apace soft, subcircular, lacking both sharp 
anterolateral border and longitudinal sulci 
anteriorly on dorsal surface. Front project- 
ing only slightly, deflected. Mxp3 with is- 
chium and merus indistinguishably fused; 
exopod present; palp 2-segmented, proximal 
longer than distal, segments articulated end- 
to-end; distal segment very short, rounded 
distally. Walking legs similar, dactyli sim- 
ple, subequal. Abdomen of 7 somites, in 
ovigerous female expanded well beyond 
bases of walking legs and mouthparts. 

Male: Unknown. 



Type species. — Nannotheres moorei, new 
species, by present designation and mono- 
typy. 

Etymology. — From the Greek, nannos, 
small, and tereo, to guard, as used in the 
name Pinnotheres. The gender is mascu- 
line. 

Host. — A bivalve mollusc of the family 
Pteriidae, Malleus candeanus (d'Orbigny). 
Waller & Macintyre (1982:490) reported 
that "In the vioihity of Carrie Bow Cay, 
Belize, specimens of Malleus candeanus 
are common from a depth of 5 m in the 
high-relief spur and groove zone seaward to 
the deepest area explored by SCUBA div- 
ing, 46 m on the steeply inclined fore-reef 
slope." 

Remarks. — Manning (1993b: 128) hsted 
four genera of pinnotherid crabs that were 
characterized by the presence of (1) simple 
dactyli on the walking legs and (2) a Mxp3 
palp composed of only two segments: the 
American genera Calyptraeotheres Cam- 
pos, 1990 and Epulotheres Manning, 
1993(a) and the Indo-west Pacific genera 
Ostracotheres Milne Edwards, 1853 and 
Xanthasia White, 1 846. In addition to being 
much smaller than species of any of these 
genera, Nannotheres differs from them as 
follows: (1) it lacks the sharp anterolateral 



312 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




1.0 mm 



Fig. L Nannotheres moorei, new genus, new species. Ovigerous female holotype, USNM 277631, frontal 
view. 



border and anterior longitudinal sulci of the 
dorsal surface of the carapace that are found 
in Calyptraeotheres; (2) it lacks the up- 
turned lateral margins and prominent, me- 
dian, mushroom-shaped tubercle found on 
the carapace of Xanthasia; (3) the Mxp3 is 
very different from that of Ostracotheres, 
in which the broad, spatulate propodus is 
much larger than the carpus; and (4) it has 
a 2-segmented palp on the Mxp3 in contrast 
to the 3-segmented palp of Epulotheres, 
which, as we will show below, was misin- 
terpreted by Manning (1993a) in his origi- 
nal account of the genus. 

Nannotheres moorei, new species 
Figs. 1, 2, 3a 

Material. — Belize, Lighthouse Reef 
[17°20'N, 87°32'W], The Blue Hole, depth 
43 m, on a dead pteriid bivalve. Malleus 
candeanus (d'Orbigny), leg. Eberhard 



Gishler: 1 ovigerous 9, cb about 1.5 mm 
(holotype, USNM 277631). 

Description. — Adult female: size small, 
carapace width of ovigerous female about 
1.5 mm. Carapace thin, membranous, ap- 
parently subcircular, regions indistinct; lat- 
eral surfaces almost vertical, lower extreme 
setose. Front depressed. Eyes very small, 
visible in dorsal view. 

Mxp3 with ischium and merus indistin- 
guishably fused, broadly ovate distally, me- 
sial margin concave proximally, strongly 
convex and weakly tuberculate distally; 
palp articulated at about midlength of inner 
distal face of ischium-merus, 2-segmented, 
proximal longer, distalmost segment short, 
rounded, terminally setose. Exopod 2-seg- 
mented. 

Chela (PI) with movable finger about as 
long as palm (measured dorsally), cutting 
edges lacking distinct teeth, cutting edge of 



VOLUME 109, NUMBER 2 



313 



fixed finger with low, long, convex, sharp 
superior flange. Palm height more than half 
dorsal length. Surfaces of palm overall 
sparsely setose, inner face with strong setae 
in lower half. 

Walking legs (P2-P5) equal right and 
left, sparsely setose, P3 and P4 lacking 
swimming setae; relative lengths, P5 < P4 
< P2 < P3; dactyli of walking legs sub- 
equal, simple, and similar; propodus of P5 
shortest of all propodi. P2 with dactylus 0.7 
times as long as propodus, latter 3.7 times 
longer than high, 1.2 times as long as car- 
pus; merus 1.7 times as long as carpus. P3 
with dactylus (missing from right P3 in Fig. 
2d) 0.7 times as long as propodus, latter 3.6 
times longer than high, 1 .2 times as long as 
carpus; merus 1.7 times as long as carpus. 
P4 with dactylus 0.7 times as long as prop- 
odus, latter 2.8 times longer than high, 1.3 
times as long as carpus; merus 1.8 times as 
long as carpus. P5 with dactylus 0.9 times 
as long as propodus, latter 2.6 times longer 
than high, 1.1 times as long as carpus; me- 
rus 1.3 times as long as carpus. 

Abdomen extending well beyond bases 
of walking legs and buccal mass, lateral 
edge of abdomen folded inward. 

Plpl biramous, both branches 2-seg- 
mented; Plp2 biramous, exopod 2-segment- 
ed, extending around outside of egg mass; 
Plp3-4 uniramous, 2-segmented. Ova large, 
17 in number, 0.35-0.5 mm in diameter. 

Size. — Unique holotype, ovigerous fe- 
male, carapace width about 1.5 mm. 

Etymology. — Named for Donald R. 
Moore, Rosenstiel School of Marine and 
Atmospheric Sciences, University of Mi- 
ami, Florida, whose keen eye spotted this 
minute crab on the edge of its host's shell 
after it had been brought on deck. 

Remarks. — Schmitt et al. (1973:9) listed 
four nominal species of Pinnotheres that 
were associated with bivalve molluscs of 
the genus Pinctada Bolten, P. margarita 
Smith, 1869, from the eastern Pacific; P. 
margaritiferae Laurie, 1906, from Ceylon; 
P. villosulus Guerin-Meneville, 1831, from 
Indonesia; and P. trichopus Tesch, 1918, 



from Indonesia. In addition to being larger, 
all of these species have three-segmented 
palps on Mxp3, so they can be separated at 
once from Nannotheres on the structure of 
the Mxp3 alone. Campos (1989) transferred 
the American P. margarita Smith to his 
new genus Tumidotheres. 

Pesta (1911) reported Pinnotheres sp. 
from a Pinctada taken in the Molucca Is- 
lands and remarked that the host had a nod- 
ular growth inside the shell that housed the 
crab. We could see no such feature in the 
host of our specimen from Belize. Barnard 
(1950) also recorded a Pinnotheres sp. from 
a Pinctada. 

Examination of the specimens of Malleus 
candeanus from Belize studied by Waller & 
Macintyre (1982) yielded no other speci- 
mens. 

Although a note accompanying the spec- 
imen suggested that it was found, dead, on 
the edge of the shell of its host, examination 
of the dried host revealed that the left chela 
of the crab remained attached to the mantle 
of the host. The crab seemed to have little 
internal tissue; the only visible muscles 
were associated with the egg-bearing pleo- 
pods. 

Our first comparison of this little crab 
was with the type species of Epulotheres 
Manning, 1993(a), E. angelae, from the Ca- 
ribbean Sea, as it, too, is small and was re- 
ported to have a two-segmented palp on 
Mxp3. Reexamination of the Mxp3 re- 
vealed that the original account was in er- 
ror, as it has a three-segmented palp (Fig. 
3^). It can be separated from Nannotheres 
on that feature alone. Epulotheres remains 
a valid genus, distinct from Pinnotheres 
S.S., for its Mxp3 has a very different shape 
from that of Pinnotheres pisum (Linnaeus, 
1767), the type species of Pinnotheres 
Bosc, 1802. In Nannotheres the fused is- 
chium-merus is much more oval, the inner 
margin lacking the angled projection found 
in the Mxp3 of Pinnotheres; the carpus and 
propodus of the Mxp3 are subequal in 
length and shaped very differently from 
those of P. pisum; and the palp is very short 



314 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Fig. 2. Nannotheres moorei, new genus, new species. Ovigerous female holotype, USNM 277631. a, Right 
chela, outer face; b. Right chela, inner face; c, Right P2, anterior face; d. Right P3, anterior face (dactylus 
missing); e, Right P4, anterior face;/ Right P5, anterior face; g. Right Plpl, anterior face; h, Right Plp2, anterior 
face; i, Right Plp3, anterior face; j. Right Plp4, anterior face. 



and inserted almost at the mid-point of the 
distal margin of the propodus in N. moorei, 
rather than being long and slender and in- 
serted basally as in P. pisum. 

We believe that this species may be the 
smallest known brachyuran crab. It is 
smaller than members of the pygmy piri- 
melid crab Sirpus Gordon, 1953(a), in 
which ovigerous females are as large as 4.5 
to 7.0 mm long in S. monodi Gordon, 
1953(b). Adults of S. gordonae Manning & 
Holthuis, 1981, are 2.5 to 3.1 mm long, and 



adults of the type species of Sirpus, S. zar- 
iquieyi Gordon, 1953(a), are 3.4 to 5.1 mm 
long (Manning & Holthuis 1981:71); ovi- 
gerous females are not known in these latter 
taxa. Nannotheres moorei also is smaller 
than all of the cryptochirid crabs mentioned 
by Kropp & Manning (1987) and Kropp 
(1989, 1990), which may be narrower but 
always are longer Ovigerous females of the 
diminutive Epulotheres angelae Manning, 
1993 are twice as wide, with carapace 
widths of 3.0 and 3.1 mm. 



VOLUME 109, NUMBER 2 



315 




Fig. 3. Outer face of MXP3. a, Nannotheres moorei, new genus, new species, ovigerous female holotype, 
USNM 277631; b, Epulotheres angelae Manning, spent female holotype, USNM 256975. 



316 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Distribution. — Known only from the 
type locality. The Blue Hole, Lighthouse 
Reef, Belize, Caribbean Sea. 

Acknowledgments 

We thank our colleague Donald R. 
Moore for bringing this little crab to our 
attention. We thank our colleagues in the 
Department of Paleobiology at the National 
Museum of Natural History, Thomas R. 
Waller for identifying the host and Warren 
Blow for helping with the identification of 
the host, examining other specimens of 
Malleus from Belize, and for providing a 
copy of Waller & Macintyre (1982). This is 
contribution no. 392 from the Smithsonian 
Marine Station at Link Port, Fort Pierce, 
Florida. Support of that facility for studies 
on the systematics of pinnotherids is grate- 
fully acknowledged. This is contribution 
no. 52 from the Laboratory for Crustacean 
Research, University of Southwestern Lou- 
isiana. 

Literature Cited 

Barnard, K. H. 1950. Descriptive catalogue of South 
African decapod Crustacea (crabs and 
shrimps). — Annals of the South African Muse- 
um 38:1-837. 

Bosc, L. A. G. 1802. Histoire naturelle des Crustaces, 
contenant leur description et leurs moeurs, avec 
figures dessinees d'apres nature 1:1-258, De- 
terville, Paris. 

Campos, E. 1989. Tumidotheres, a new genus for Pin- 
notheres margarita Smith, 1869, and Pinnoth- 
eres maculatus Say, 1818 (Brachyura: Pinnoth- 
eridae). — Journal of Crustacean Biology 9:672- 
679. 

. 1990. Calyptraeotheres, a new genus of Pin- 

notheridae for the limpet crab Fabia granti 
Glassell, 1933 (Crustacea, Brachyura). — Pro- 
ceedings of the Biological Society of Washing- 
ton 103:364-371. 

Gordon, I. 1953a. On a new crab from Cadaques on 
the north east coast of Spain (Sirpus zariquieyi 
n. gen and sp.). — Eos, Revista Espanola de En- 
tomologia 28(4):303-314. 

. 1953b. On S/r/7Mj, a genus of pigmy cancroid 

crabs. — Bulletin of the British Museum (Natu- 
ral History), Zoology 2(3):43-65. 

Guerin-Meneville, F E. 1829-1844. Iconographie du 
regne animal de G. Cuvier, ou representation 



d'apres nature de I'une des especes les plus re- 
marquables, et souvent non figurees, de chaque 
genre d'animaux, avec un text descriptif mis au 
courant de la science: ouvrage pouvant servir 
d' atlas a tous les traites de zoologie. 450 plates 
in 45 livraisons. Crustacea: 36 pis, 48 pp. Paris. 

Kropp, R. K. 1989. A revision of the Pacific species 
of gall crabs, genus Opecarcinus (Crustacea: 
Cryptochiridae). — Bulletin of Marine Science 
45:98-129. 

. 1990. Revision of the genera of gall crabs 

(Crustacea: Cryptochiridae) occurring in the Pa- 
cific Ocean. — Bulletin of Marine Science 44(4): 
417-448. 

, & R. B. Manning. 1987. The Atlantic gall 



crabs, family Cryptochiridae (Crustacea: De- 
capoda: Brachyura). — Smithsonian Contribu- 
tions to Zoology 462:21 pp. 

Laurie, R. D. 1906. Report on the Brachyura collected 
by Professor Herdman, at Ceylon, in 1902. In 
W. A. Herdman, ed.. Report to the government 
of Ceylon on the pearl oyster fisheries of the 
Gulf of Manaar, 5 (Supplemental Reports, no. 
40):349-432, pis. 1, 2. 

Linnaeus, C. 1767. Systema naturae per regna tria na- 
turae, secundum classes, ordines, genera, spe- 
cies, cum characteribus, differentiis, synonymis, 
locis. Twelfth edition. Stockholm, 1(2):533- 
1327. 

Manning, R. B. 1993a. Epulotheres angelae, new ge- 
nus, new species, a pinnotherid crab from the 
Caribbean Sea (Decapoda: Pinnotheridae). — 
Journal of Crustacean Biology 13:801-804. 

. 1993b. West African pinnotherid crabs, fam- 
ily Pinnotherinae (Crustacea, Decapoda, 
Brachyura). — Bulletin du Museum National 
d'Histoire Naturelle, Paris, serie 4, 15(Sec. A, 
nos. 1-4): 125-177. 

, & L. B. Holthuis. 1981. West African 



brachyuran crabs. — Smithsonian Contributions 
to Zoology 306:379 pp. 

Milne Edwards, H. 1853. Memoire sur la famille des 
Ocypodiens, suite. — Annales des Sciences Na- 
turelles, serie 3 (Zoologie) 20:163-228. 

Pesta, O. 1911. Decapoda Brachyura aus Samoa (Un- 
ter Beriicksichtigung der Sammlungen des k.k. 
Naturhistorischen Hofmuseums in Wien). Crus- 
tacea, I. Teil. In K. Rechinger, ed., Botanische 
und zoologische Ergebnisse einer wissenschaf- 
ten Forschungsreise nach den Samoainseln, dem 
Neuguinea Archipel und den Salomoninseln 
Marz bis Dezember 1905. — Denkschriften der 
Mathematisch-naturwissenschaftlichen Klasse 
der Kaiserlichen Akademie der Wissenschaften, 
Wien 88:36-55, pi. 3. 

Schmitt, W L., J. C. McCain, & E. S. Davidson. 1973. 
Fam. Pinnotheridae: Brachyura I: Decapoda I. 
Pp. 1-160 in H.-E. Gruner & L. B. Holthuis, 



VOLUME 109, NUMBER 2 



317 



eds., Crustaceorum Catalogus, 3. W. Junk, Den 
Haag, The Netherlands. 

Smith, S. I. 1869. [Untitled footnotes]. Pp. 245-250 
in A. E. Verrill, ed.. On the parasitic habits of 
Crustacea. — American Naturalist 3:239-250. 

Tesch, J. J. 1918. Goneplacidae and Pinnotheridae. 
The Decapoda Brachyura of the Siboga Expe- 
dition, II.— Siboga-Expeditie, 39(c'): 149-295, 
pis. 7-18. 

Waller, T. R., & I. G. Macintyre. 182. Larval settle- 



ment behavior and shell morphology of Malleus 
candeanus (d'Orbigny) (MoUusca: Bivalvia). 
Pp. 489-497 in K. Riitzler & I. G. Macintyre, 
eds., The Atlantic barrier reef ecosystem at Car- 
rie Bow Cay, Belize, I. Structure and commu- 
nities. — Smithsonian Contributions to the Ma- 
rine Sciences 12:539 pp. 
White, A. 1846. Notes on four new genera of Crus- 
tacea. — Annals and Magazine of Natural His- 
tory 18:176-178. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 

109(2):318-325. 1996. 

New species of Macrogynoplax (Insecta: Plecoptera: Perlidae) from 

Peru and Guyana 

Bill P. Stark 

Department of Biology, Mississippi College, 
Clinton, Mississippi 39058, U.S.A. 

Abstract. — Macrogynoplax truncata, new species, and M. yupanqui, new 
species, are described from specimens collected in Peru, and M. flinti, new 
species, and M. kanuku, new species, are described from specimens collected 
in Guyana. Holotype males are designated for each species and a revised key 
is provided for adults of the nine species known for the genus. 



Macrogynoplax currently includes five 
uncommon species, each apparently endem- 
ic to remote regions of eastern South Amer- 
ica (Zwick 1973, Froehlich 1984, Stark & 
Zwick 1989). Two species (M. neblina 
Stark, in Stark & Zwick 1989:248 and M. 
spangleri Stark, in Stark & Zwick 1989:251) 
are known from Cerro de la Neblina in 
southern Venezuela and the others (M. ge- 
ijskesii Zwick, in Stark & Zwick 1989:253, 
M. guayanensis Enderlein, 1909:403, andM. 
veneranda Froehlich, 1984:39) are known 
from a few specimens from Suriname, Guy- 
ana, or southeastern Brazil. In this study I 
report the occurence of two undescribed Ma- 
crogynoplax species in Peru and an addi- 
tional undescribed pair in southern Guyana. 
Holotypes are deposited in the National Mu- 
seum of Natural History (USNM). 

Macrogynoplax yupanqui, new species 
Figs. 1-4 

Adult habitus. — Biocellate. White in al- 
cohol, possibly green in life. Unpigmented 
lobes along inner margins of compound 
eyes. 

Male. — Forewing length 12 mm. Para- 
procts prominently spined on anteroapical 
margin (Fig. 2). Sternum 9 with a pair of 
basolateral knobs; hammer inflated, bulb- 
like (Fig. 1). Aedeagal tube covered with 
an outer skirtlike membranous sheath 



armed with fine transparent spines; tube 
with an inner, partially exposed sclerite; 
posteromedian margin of sclerite notched, 
and sclerotized lateral margins convergent 
apically. Short, membanous, apical aedea- 
gal section bilobed on ventromedian sur- 
face; apex sparsely clothed with fine trans- 
parent setae (Figs. 3-4). 

Female. — Unknown. 

Nymph. — Unknown. 

Etymology. — This species honors Tupa 
Inca, or Yupanqui, tenth ruler in the Inca 
dynasty. 

Types. — Holotype S from Rio Tambo- 
pata, 30 km SW Maldonado, 290 m, Madre 
de Dios, Peru, 21-25 Oct 1979, J. B. Hepp- 
ner (USNM). 

Macrogynoplax truncata, new species 
Figs. 5-9 

Adult habitus. — Biocellate. White in al- 
cohol, possibly green in life. Unpigmented 
lobes along inner margins of compound 
eyes. 

Male. — Forewing length 10 mm. Para- 
procts small, anteroapical spine minute 
(Fig. 7). Hammer rectangular (Fig. 5). Ae- 
deagal tube enclosed by an outer skirtlike 
membranous sheath armed with fine trans- 
parent spines; inner sclerite partially ex- 
posed apically; sclerotized lateral margins 
of sclerite convergent apically. Short, mem- 



VOLUME 109, NUMBER 2 



319 






Figs. 1-4. Macrogynoplax yupanqui male genitalia. 1. Sternum 9; 2. Paraproct, lateral; 3. Aedeagal tube, 
ventral; 4. Aedeagal tube, lateral. 



320 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




S?lJ;-,-.'4-«?.?^>r.- ■/ -1 !■ / • ■•/,■■ < ■ •.•■.:.;%.^.i..,:,^- 

•l:n;-/, ^. ;'r''-''' '' ■'• • ■''/■■ '" ': '■■ '■'<'--^'-,- 
-.■■^'■'.■n ;■■■.■' ;■•' -I- -, ; /• !■'■':'■> t 

{■■■■■■■ '•■■'. .1 ■ I ■■■,■' ■■ -■'/•(■ 





Figs. 5—9. Macrogynoplax truncata male and female genitalia. 5. Male sternum 9; 6. Female sterna 8 and 
9; 7. Paraproct, lateral; 8. Aedeagal tube, ventral; 9. Aedeagal tube, lateral. 



branous, apical aedeagal section sparsely Female. — Forewing length 12 mm. Su- 

clothed with fine transparent setae; apex bgenital plate covers most of sternum 9; 

finely divided into a pair of small membra- truncate posterior margin heavily sclero- 

nous lobes (Figs. 8-9). tized. Sternum 9 lateral bars short; median 



VOLUME 109, NUMBER 2 



321 



field with a narrow longitudinal sclerite 
(Fig. 6). 

Nymph. — Unknown. 

Etymology. — The species name refers to 
the truncate posterior margin of the female 
subgenital plate. 

Types. — Holotype S and paratype ? 
from Rio Nanay, 25 km SW Iquitos, 120 
m, Callicebus Res. Station, Loreto, Peru, 
10-17 Jan 1980, J. B. Heppner (USNM). 

Macrogynoplax flinti, new species 
Figs. 10-13 

Adult habitus. — Biocellate. Pinned spec- 
imens white with slight green tint in wing 
veins; possibly green in life. Unpigmented 
lobe along inner margins of compound 
eyes. 

Male. — Forewing length 9 mm. Para- 
procts small, anteroapical spine minute. 
Hammer outline triangular; sternum 9 with 
a pair of basolateral knobs (Fig. 10). Ae- 
deagal tube partially enclosed by a mem- 
branous sheath armed with fine transparent 
spines; tube armed with a prominent ven- 
trobasal pair of hooks and an anteapical 
shelf like process; tube apex with a pair of 
small membranous lobes, sparsely clothed 
with fine setae, and a long slender process 
(Figs. 11-13). 

Fema le. — Unknown . 

Nymph. — Unknown. 

Etymology. — The patronym honors O. S. 
Flint, Jr., collector of the type series, for his 
outstanding contributions to our knowledge 
of Neotropical aquatic entomology. 

Types. — Holotype 6 and paratype 6 
from Aramatani Creek, Dubulay Ranch, 
Guyana, 5°39.4'N, 57°55.5'W, 15-18 Apr 
1995, O. S. Fhnt (USNM). 

Habitat. — Aramatani Creek is a shallow 
stream about two meters in width flowing 
through an area of wet rain forest. The sub- 
strate is mixed sand and organic debris and 
the normally clear water develops a tea-col- 
ored stain from runoff following periods of 
heavy rainfall (O. S. Flint, pers. comm.). 



Macrogynoplax kanuku, new species 
Figs. 14-18 

Adult habitus. — Biocellate. Pinned spec- 
imens white with slight green tint in wing 
veins; possibly green in life. Unpigmented 
lobes along inner margins of compound 
eyes. 

Male. — Forewing length 9 mm. Para- 
procts small, anteroapical spine minute. 
Hammer outline elongate oval, orientation 
vertical on upturned apex of sternum 9; ba- 
solateral knobs present on sternum 9 (Fig. 
14). Aedeagal tube partially enclosed by a 
membranous sheath armed with fine trans- 
parent spines; tube armed with a prominent 
ventrobasal pair of hooks; distal section a 
simple, partially sclerotized cylinder with a 
pair of small apical lobes, sparsely clothed 
with fine setae (Figs. 16-18). 

Female. — Forewing length 12 mm. Sub- 
genital plate covers about half of sternum 
9; posterior margin rounded and notched. 
Sternum 9 lateral bars narrowly separated 
on midline; median field with a slender pale 
sclerite (Fig. 15). 

Nymph. — Unknown. 

Etymology. — The species name is based 
on the Kanuku Mountains of southern Guy- 
ana. 

Types. — Holotype S and paratype 2 
from Kumu River, Kanuku Mountains, 
Guyana, 3°15.9'N, 59°43.5'W, 28-30 Apr 
1995, O. S. Flint (USNM). 

Habitat. — Kumu River at the collecting 
site is 2-3 meters in width and tumbles 
down a heavily forested slope. Substrate 
varies from sand to boulders and there are 
numerous small springs, falls and plunge 
basins (O. S. Flint, pers. comm.). 

Discussion. — Macrogynoplax truncata 
and M. yupanqui are similar to M. spangleri 
in aedeagal structure and in shape of the 
male 9th sternum. M. truncata males are 
easily distinguished from all known Macro- 
gynoplax by hammer shape (Fig. 5) and by 
the apical aedeagal shape (Figs. 8-9). Fe- 
males of M. truncata are the only known 
species with a truncate subgenital plate 



322 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Figs. 10-13. Macrogynoplax flinti male genitalia. 10. Sternum 9; 11. Aedeagal tube, lateral; 12. Aedeagal 
tube, ventral; 13. Aedeagal tube, dorsal. 



VOLUME 109, NUMBER 2 



323 



m^ 












•if- 



-|-: ••/. 



■c >■ 



pi::; fec;^ii Pi 






:/"•/-/: 






■/••: 



■^^■? 



■.V?V 



;v;r 



•.'■■/■. 



y. ■ (•' 



■(. -.r 






14 





17 




Figs. 14-18. Macrogynoplax kanuku male and female genitalia. 14. Male sternum 9; 15. Female sterna 
and 9; 16. Aedeagal tube, ventral; 17. Aedeagal tube, lateral; 18. Aedeagal tube, dorsal. 



(Fig. 6). Macrogynoplax yupanqui males 
key to M. spangleri in Stark & Zwick 
(1989) but they are distinguished on the ba- 
sis of the strong paraproct spine (Fig. 2) and 
by the paired ventroapical aedeagal lobes 
(Fig. 3). Macrogynoplax flinti and M. ka- 



nuku share prominent sclerotized aedeagal 
hooks with M. geijskesii (Figs. 12, 16), but 
these two species seem more closely related 
to each other than either is to M. geijskesii. 
Macrogynoplax flinti can be distinguished 
on the basis of the triangular hammer (Fig. 



324 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



10) and the shelflike ventroapical aedeagal 
process (Fig. 12), but M. kanuku can prob- 
ably be reliably separated from M. geijskesii 
only on the basis of the complexly lobed 
apical aedeagal section in the latter species. 
The subgenital plate of female M. kanuku 
is similar to that of M. spangleri, but per- 
haps is not as long, and the pale mesal 
sclerite of sternum 9 (Fig. 15), should dis- 
tinguish females of these species. The fol- 
lowing key to adult specimens is modified 
from Stark & Zwick (1989). 

Males 

1 . Large subterminal sclerotized hooks 
on venter of aedeagal tube (Figs. 12, 
16) 2 

- Sclerotized aedeagal hooks absent 
(Fig. 3) 4 

2. Hammer oriented vertically on up- 
turned margin of sternum 9, outline 
oval (Fig. 14) 3 

- Hammer oriented horizontally, apex 
of sternum 9 not upturned, outline 
triangular (Fig. 10) flinti 

3. Aedeagal apex a simple cylinder 
with a pair of distal, sparsely hirsute 
lobes (Figs. 14-18) kanuku 

- Aedeagal apex complexly lobed 

geijskesii 

4. Hammer outline rectangular (Fig. 5) 
truncata 

- Hammer outline circular (Fig. 1) . . 5 

5. Sternum 9 apical plate strongly con- 
stricted at base, sides distinctly con- 
vex 6 

- Sternum 9 apical plate slightly or not 
at all constricted at base, sides par- 
allel to slightly convex (Fig. 1) . . . . 7 

6. Aedeagal apex expanded laterally, 
sclerites almost joined proximal to 
apex veneranda 

- Aedeagal apex gradually narrowed, 
sclerites separated except for apices 
which almost touch neblina 

7. Aedeagal apex with paired dorsal 
lobes (Fig. 3); paraproct spine 
curved downward (Fig. 2) . . . yupanqui 



— Aedeagal apex without dorsal lobes; 
paraproct spine straight (Fig. 7) 
spangleri 



Females 

(M. geijskesii, M. flinti and M. yupanqui 
unknown) 

1. Subgenital plate truncate (Fig. 6) 
truncata 

- Subgenital plate notched or rounded 
2 

2. Subgenital plate extending beyond 
posterior margin of sternum 9 

guyanensis 

- Subgenital plate not reaching be- 
yond posterior margin of sternum 9 
3 

3. Lateral bars of sternum 9 fused mesal- 

ly under subgenital plate neblina 

- Lateral bars of sternum 9 separated 
mesally 4 

4. Subgenital plate almost reaching 
posterior margin of sternum 9; lat- 
eral bars narrowly separated 

spangleri 

- Subgenital plate reaching slightly 
beyond midpoint of sternum 9; lat- 
eral bars widely or narrowly sepa- 
rated 5 

5. Lateral bars of sternum 9 widely 

separated veneranda 

Lateral bars of sternum 9 narrowly 
separated kanuku 



Acknowledgments 

I am grateful to N. Adams and O. S. Flint 
(National Museum of Natural History) for 
arranging the loan of material used in this 
study. O. S. Flint also provided field notes 
on the Guyana collecting sites. 

Literature Cited 

Enderlein, G. 1909. Klassifikation der Plecopteren, 
sowie Diagnosen neuer Gattungen und Arten. — 
Zoologischer Anzeiger 34:385-419. 



VOLUME 109, NUMBER 2 325 

Froehlich, C. 1984. Brazilian Plecoptera 3. Macro- coptera:Perlidae). — Aquatic Insects 11:247- 

gynoplax veneranda Sp. n. (Perlidae: Acroneu- 255. 

riinae). — Annales de Limnologie 20:39-42. Zwick, P. 1973. Die Plecopteren-Arten Enderleins (In- 
Stark, B. P., & P. Zwick. 1989. New species of Ma- secta); Revision derTypen. — Annales Zoologici 

crogynoplax from Venezuela and Surinam (Pie- 30:471-507. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2): 326-348. 1996. 

Australian beach flies (Diptera: Canacidae) 

Wayne N. Mathis 

Department of Entomology, National Museum of Natural History, MRC 169, 
Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — The Australian fauna of beach flies (8 genera and 14 species), 
excluding Zaleinae, is reviewed, including description of seven new species as 
follows (type locality indicated in parentheses): Nocticanace australina (NSW: 
Lord Howe Island, Roach Island Beach), Procanace mcalpinei (NSW: Karuah), 
Chaetocanace flavipes (WA: West Kimberley, 4 km SSW of Cape Bertholet), 
Chaetocanace koongarra (WA: Koongarra, 15 km E of Mt. Cahill), Chaeto- 
canace longicauda (NT: 35 km W Jabiru, South Alligator River area), Dynom- 
iella australica (TAS: Squeaking Point near Port Sorell), and Xanthocanace 
collessi (WA: West Kimberley, 5 km SSW of Cape Bertholet). Procanace Hen- 
del and Dynomiella Giordani Soika are reported for the first time from Aus- 
tralia. An undescribed genus and species, represented by a single female, is 
also reported. 



The Australian fauna of the family Can- 
acidae, more commonly known as beach or 
surf flies, has never been treated compre- 
hensively. Only the subfamily Zaleinae, 
which is questionably a canacid, has been 
considered recently (McAlpine 1982, 
1985). Although relatively few species oc- 
curring in Australia have been described, 
nine species in six genera (Mathis 1989a, 
Colless & McAlpine 1991), additional taxa 
have been discovered recently, and descrip- 
tion of these, within the context of a faunal 
review, is the primary purpose of this paper 

Three of the nine described species are 
in the genus Zalea McAlpine. This enig- 
matic genus has been segregated as the sub- 
family Zaleinae (McAlpine 1982, 1985) but 
is perhaps better placed in the family Teth- 
inidae (Mathis 1992, Freidberg 1995). Za- 
lea, which was recently revised (McAlpine 
1982, 1985), is not included here. 

The new species described herein mostly 
represent recent discoveries but also clarify 
the status of an unidentified species of Noc- 
ticanace Malloch that was reported previ- 
ously from Australia (Colless & McAlpine 
1991). Two of the new species are repre- 



sentatives of Procanace Hendel and Dy- 
nomiella Giordani Soika, both genera pre- 
viously unknown from Australia. The Aus- 
tralian species of Dynomiella is of particu- 
lar interest, being the first record of this 
genus outside of Africa and representing a 
significant range extension. Four of the oth- 
er six new species belong in genera previ- 
ously known from Australia {Chaetocanace 
Hendel and Xanthocanace Hendel), and the 
remaining species, perhaps representing a 
new genus, is included in keys and discus- 
sions but is not described, being known 
from a single female. 

The more inclusive purpose of this paper 
is to document comprehensively the Aus- 
tralian fauna (excluding the subfamily Zal- 
einae), including keys to genera and species 
known from the region. Illustrations, partic- 
ularly of characters of the external male ter- 
minalia to aid in species identification, and 
locality data to understand better the distri- 
bution of the included taxa are also provid- 
ed. This effort is intended to foster further 
interest in the natural history and phyloge- 
ny of the family, which is almost exclu- 



VOLUME 109, NUMBER 2 



327 



sively associated with maritime beaches in 
temperate and tropical regions of the world. 

Methods and Materials 

For each genus and species reviewed, a 
synonymy, the known distribution, includ- 
ing detailed information on the type local- 
ity, the depository of the primary types, 
specific locality data for Australia, the gen- 
eralized distribution, a diagnosis, and a re- 
marks section, as appropriate, are provided. 
In the synonymies, only literature that is 
pertinent to the Australasian Region or a 
taxon's nomenclatural history is cited. For 
new species, a more complete description is 
provided, in addition to characters included 
in diagnoses of appropriate species groups 
and genera. The species groups that are rec- 
ognized for the genera Nocticanace and 
Procanace are not necessarily monophylet- 
ic assemblages and are used primarily to 
divide conveniently the genera into man- 
ageable units and to facilitate their identi- 
fication. Their characterization and the spe- 
cies composition may change as a result of 
future revisionary and phylogenetic studies. 
The descriptive format for the new species 
follows Mathis & Wirth (1979) and Mathis 
(1982, 1988). More details concerning the 
morphology and higher classification of 
Canacidae are found in Mathis (1982) and 
Wirth (1987). 

Two venational ratios are used in the de- 
scriptions. Costal vein ratio: The straight 
line distance between the apices of vein 
R2+3 and R4+5/distance between the apices 
of veins Rj and Ra+s- M vein ratio: The 
straight line distance along vein M between 
crossveins (dm-cu and r-m)/distance apicad 
of dm-cu. 

Specimens, holotypes in particular, are 
housed in the following institutions (acro- 
nyms are used in the descriptive portion of 
this paper). 

AM Australian Museum, Sydney (Dr. 
David K. Mc Alpine). 
ANIC The Australian National Insect 
Collection, CSIRO, Division of 



Entomology, Canberra (Dr. Peter 
S. Cranston). 

BMNH The Natural History Museum 
(formerly the British Museum 
(Natural History)), London, En- 
gland (Dr. Brian Pitkin). 
UQIC University of Queensland Insect 
Collection, Brisbane (Ms Mar- 
garet A. Schneider). 

USNM former United States National 
Museum, collections in the Na- 
tional Museum of Natural Histo- 
ry, Smithsonian Institution, 
Washington, D.C. 
ZIL Zoological Institute, University 
of Lund, Sweden (Dr. Roy Dan- 
ielsson). 

Family Canacidae Jones 

Key to Australian Genera of Canacidae 

1. Lateroclinate fronto-orbital setae 3; 
katepisternal seta usually present 
(lacking in the grisescens group of 
Procanace); lamella of 2 terminalia 
bearing 2 moderately large setae, 
one apical the other subapical, each 
rather bluntly rounded (subfamily 
Nocticanacinae) 2 

- Lateroclinate fronto-orbital setae ei- 
ther 4 or more, or 3 and with kate- 
pisternal seta lacking (subfamily 
Canacinae) 3 

2. Intrafrontal setae absent, although 
anterior % of frons occasionally with 
scattered setulae . . . .Procanace Hendel 

- One or 2 intrafrontal setae in addi- 
tion to any other setulae 

Nocticanace Malloch 

3. Lamella of 2 reduced and lacking 
any large setae new genus 

- Lamella of 2 terminalia large with 
1 large, apical seta, this usually 
acutely pointed 4 

4. Anterior notopleural seta present; 
lateral scutellar setae usually 2 pairs 
5 



328 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



- Anterior notopleural seta absent; lat- 
eral scutellar setae 1 pair 7 

5. Vein M with last section arcuate; 
mesofrons uniformly and densely 
setulose; 4-6 fronto-orbital setae; 
setae generally pale 

Xanthocanace Hendel 

- Vein M with last section more or 
less straight, not distinctly arcuate; 
mesofrons with bare areas, not 
densely setulose; 4 fronto-orbital se- 
tae; setae generally dark colored . . 6 

6. Arista bearing 2 rows of hairs, these 
extended from base to apex, 1 row 
dorsal, the other ventral; postocellar 
setae conspicuously smaller than 
ocellar setae and with proclinate and 
slightly divergent orientation (the al- 
biceps group) Isocanace Mathis 

- Arista with at least apical Vs bare; 
postocellar setae subequal in size 
and with similar orientation as ocel- 
lar setae . . . Dynomiella Giordani Soika 

7. Mesofrons well sclerotized, distinct 
from membranous-appearing para- 
frons; anteroclinate genal seta, large, 
black; upturned genal setae lacking; 
arista with 2 rows of setulae extend- 
ed to apex; postpronotum setulose 

Chaetocanace Hendel 

- Mesofrons and parafrons not distinct 
from each other except by color in 
some species, both appearing mem- 
branous although usually microto- 
mentose; anteroclinate genal seta 
small, pale; upturned genal setae 2; 
arista with apical % to Vi bare, style- 
like; postpronotum bare of setulae 
Trichocanace Wirth 



Subfamily Nocticanacinae Mathis 
Genus Nocticanace Malloch 

Nocticanace Malloch, 1933:4. Type spe- 
cies: A^. peculiaris Malloch, by original 
designation. — Mathis, 1989b:594 [key to 
species groups]; 1992:8-10 [world cata- 
log]. 



Diagnosis. — Small to medium-sized 
beach flies, length 1.8 to 3.7 mm; general 
coloration brown to grayish black. 

Head: Intrafrontal setae 1 pair; postocel- 
lar setae either absent or much reduced, less 
than Va length of ocellar setae; ocelli form- 
ing an isosceles triangle, distance between 
posterior ocelli greater than that between ei- 
ther posterior ocellus and the anterior ocel- 
lus. Lower facial margin sinuous; clypeus 
low, width subequal to length of antenna. 
Two large upturned genal setae; anterocli- 
nate genal seta moderately well developed, 
at least V2 length of larger upturned genal 
setae. Palpus grayish black, bearing 1 to 
several long setae, each seta 2 to 3 times as 
long as greatest width of palpus. 

Thorax: Anepisternum with scattered 
setulae; proepisternal seta absent; katepis- 
ternal seta present, well developed. Legs 
entirely grayish black; forefemur bearing 
4-6 long, evenly spaced setae along pos- 
teroventral margin, length of setae at least 
equal to and usually greater than width of 
femur 

Discussion. — Nocticanace, now with 35 
species (Mathis 1991), has more species 
than any other genus of the family. These 
have been divided into five species groups 
(Mathis 1989b, Mathis & Freidberg 1991), 
mostly to facilitate their identification. The 
new Australian species described below is 
part of the pacifica group, the largest group 
in the genus with 22 species. This genus 
probably occurs on mainland Australia in 
addition to Lord Howe Island. 

The pacifica Group 

Diagnosis. — General coloration dark, 
grayish brown to grayish black. Thorax: 
Acrostichal setulae absent; apical scutellar 
setae distinctly upturned; anterior noto- 
pleural seta absent; proepisternal seta(e) 
present; anepisternum with scattered setu- 
lae; katepisternal seta present. Legs entirely 
dark, grayish brown to black; forefemur 
with 4-6 long and evenly spaced setae 
along posteroventral margin, length greater 



VOLUME 109, NUMBER 2 



329 




Figs. 1-2. External male terminalia of Nocticana- 
ce australina: 1, epandrium, cercus, and surstylus, lat- 
eral view; 2, surstylus, ventrolateral view. Scale = 0.1 
mm. 



than width of femur; midfemur of male 
without combhke row of setae; hindtibia 
lacking spinelike setae apically. Wing with 
length of apical section of vein CuAj long, 
about twice length of cross vein dm-cu; M 
vein ratio 0.45-0.5. 

Nocticanace australina, new species 
Figs. 1-2 

Nocticanace species, Colless & McAlpine, 
1991:779. 

Diagnosis. — This species is distin- 
guished from congeners, especially of the 
pacifica group, by its dark coloration (gray- 
ish brown to charcoal) and by the napiform 
shape of the surstylus in lateral view. 

Description. — Small to moderately small 
beach flies, length 1 .9 to 2.4 mm; generally 
dark colored, densely microtomentose, ap- 
pearing dull, grayish brown to charcoal 
black. 

Head: Frons grayish black. Antenna 
black. Face whitish gray to grayish black 



depending on angle of view; gena similar 
in color to face. 

Thorax: Mesonotum brown to brownish 
black; pleural areas gray to charcoal gray. 
Wing lightly infuscate, faintly brownish 
black; costal vein ratio 0.17; M vein ratio 
0.5. Legs uniformly and entirely grayish 
black. 

Abdomen: External male terminalia: sur- 
stylus in lateral view (Figs. 1-2) napiform, 
anterior and posterior margin shallowly 
rounded, ventral margin produced to form 
a narrow ventromedial process that is 
curved anteroventrally; posterior margin of 
surstylus bearing fringe of long setulae, 
these oriented medially. 

Type material. — The holotype male is la- 
beled "[Australia. New South Wales:] 
Roach Isl. Beach[,] Lord Howe Isl[and].[,] 
23 Feb 1988[,] H. J. de S. Disney[,] Rock 
pools/Austr. Mus. Collection." Paratypes 
are as follows: Australia. New South Wales: 
Lord Howe Island. Old Gulch, Dec 1972, 
Z. Liepa (1 S\ ANIC); Roach Island Beach 
(rock pools), 23 Feb 1988, H. J. de S. Dis- 
ney (7 d; AM, USNM); Salmon Beach, 
Dec 1972, Z. Liepa (1 2; ANIC). The ho- 
lotype is double mounted (glued to a paper 
point), is in excellent condition, and is de- 
posited in the ANIC. 

Distribution. — Australasian: Eastern 
Austraha (NSW). 

Etymology. — The specific epithet, aus- 
tralina, refers to the continent where this 
species occurs. 

Remarks. — Colless & McAlpine (1991) 
first listed this species, although then un- 
described, from Australia. 

Genus Procanace Hendel 

Procanace Hendel, 1913:93. Type species: 
Procanace grisescens Hendel, by original 
designation.— Wirth, 1951:253-259 [re- 
vision].— Delfinado, 1970:527-531 [re- 
vision of species from New Guinea]. — 
Delfinado & Wirth, 1977:392 [Oriental 
catalog].— Hardy & Delfinado, 1980: 
388-406 [revision of Hawaiian spe- 



330 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



cies].— Mathis, 1989a:669-670 [Austral- 
asian/Oceanian catalog]; 1989b:603-607 
[key to species groups]; 1992:10-12 
[world catalog]. 

Diagnosis. — General coloration whitish 
gray, olivaceous, to blackish brown. 

Head: Intrafrontal setae absent, but with 
a few setulae inserted anteriorly; fronto-or- 
bital setae 3; ocelli forming equilateral or 
isosceles triangle, if isosceles, the greater 
distance is between posterior ocelli. Arista 
pubescent over entire length. Two large up- 
turned genal setae; anteroclinate genal seta 
moderately well developed. Palpus not 
bearing long setae. Epistomal margin, in 
lateral view, more or less horizontal. 

Thorax: Acrostichal setae, especially a 
prescutellar pair of large setae, usually lack- 
ing (setulae present in species of the wil- 
liamsi group); scutellar disc lacking setae 
(1-2 pairs of scutellar disc setulae occur in 
P. nakazatoi Miyagi of the williamsi 
group); 2 pairs of marginal scutellar setae, 
apical pair not upturned; anterior and pos- 
terior notopleural setae present, length of 
both subequal; anepisternum with scattered 
setulae. Katepisternal setal usually present 
(lacking in species of the grisescens group). 
Hindtibia lacking spinelike setae apically. 

Abdomen: Male genitalia as follows: 
epandrium in posterior view wider than 
high; cerci reduced, poorly sclerotized; sur- 
stylus with an anterior and a posterior lobe, 
the latter larger, sometimes markedly so and 
shape unique to species. 

Discussion. — Procanace, now with 30 
species, is the second largest genus in the 
family. Mathis (1989b) divided the genus 
into five species groups, largely to expedite 
identification of the various species. Some 
of the groups are monophyletic, but others 
may not be. The new Australian species is 
in the fulva group, which is diagnosed be- 
low. 

ThQ fulva Group 

Diagnosis. — General coloration whitish 
olivaceous to brown. Head: Postocellar se- 




Figs. 3-4. External male terminalia of Procanace 
mcalpinei: 3, epandrium, and surstylus, lateral view; 
4, surstylus, posteroventral view. Scale = 0.1 mm. 



tae either absent or much reduced; ocelli 
forming equilateral triangle. Palpus yellow. 
Clypeus low, height VS to Va eye height. 
Thorax: Acrostichal setulae absent; proepi- 
stemal seta(e) usually present; anepister- 
num with scattered setulae; katepisternal 
seta present. Femora and tibiae grayish, tar- 
si yellowish; forefemur lacking 4-6 long 
and evenly spaced setae along posteroven- 
tral margin, length greater than width of fe- 
mur; midfemur of male bearing comblike 
sparse row of setae. Wing with length of 
apical section of vein CuAi short, subequal 
to or slightly longer than crossvein dm-cu; 
M vein ratio 0.75 to 0.8. 

Remarks. — The fulva group, now with 
nine species, occurs along the western mar- 
gin of the Pacific Ocean (eastern Palearctic, 
Oriental, and Australasian regions). 

Procanace mcalpinei, new species 
Figs. 3-4 

Diagnosis. — This species is distin- 
guished from congeners by the shape of the 



VOLUME 109, NUMBER 2 



331 



surstylus, which in lateral view is irregular- 
ly ovate. The posterior margin of the sur- 
stylus bears a median tooth and short emar- 
gination just dorsad of tooth. 

Description. — Moderately small beach 
flies, length 2.0 mm; generally grayish 
brown to brown, generally dull, microto- 
mentose. 

Head: Mesofrons brown, similar to 
mesonotum; parafrons more blackish 
brown; postocellar setae reduced, about Vi 
length of ocellar setae, proclinate slightly 
divergent. Scape and pedicel grayish black; 
1 St flagellomere brownish yellow with faint 
blackish tinges. Face grayish white; gena 
low, about Vi eye height, concolorous with 
face anteriorly, yellowish along anteroven- 
tral margin of eye, becoming slightly grayer 
posteriorly. 

Thorax: Mesonotum light brown to faint- 
ly olive brown, dull; pleural areas grayer. 
Wing lightly infumate, faintly brownish; 
costal vein ratio 0.18; M vein ratio 0.8. 
Legs brownish yellow; femora thinly in- 
vested with whitish microtomentum. 

Abdomen: Dorsum concolorous with 
mesonotum. External male terminalia as 
follows: epandrium bearing numerous long 
setae; surstylus in lateral view (Fig. 3-4) 
irregularly ovate, median surface bearing 
dorsoventral row of stout setulae, posterior 
margin (Fig. 4) with short emargination at 
dorsal Va and an edentate projection medi- 
ally. 

Type material. — The holotype male is la- 
beled "[AUSTRALIA.] Karuah, N.S.W. 
23.xii.1968 [23 dec 1968] inlet: beach L C. 
Yeo." The holotype is double mounted, is 
in good condition (right antenna missing, 
abdomen removed and dissected, structures 
in an attached microvial), and is deposited 
in UQIC. 

Distribution. — This species is known 
only from the type locality. 

Etymology. — The specific epithet, mcal- 
pinei, is a genitive patronym to recognize 
and honor the contributions of David K. 
McAlpine to the study of acalyptrate Dip- 
tera, the Australian fauna in particular. 



Subfamily Canacinae Jones 

Tribe Dynomiellini Mathis 

Genus Chaetocanace Hendel 

Chaetocanace Hendel, 1914:98. Type spe- 
cies: Canace biseta Hendel, by original 
designation and monotypy. — Malloch, 
1924:333 [generic key].— Curran, 1934: 
357 [generic key]. — Wirth, 1951:265 [re- 
view]. — Mathis, 1982:7-9 [review]; 
1992:5 [world catalog]. 

Diagnosis. — Resembling Isocanace but 
differing from it and other genera by the 
following combination of characters. 

Head: Mesofrons distinct from parafrons, 
shinier, less microtomentose, with 4-6 
large, lateral, generally proclinate setae, 
middle area bare; postocellar setae subequal 
to ocellar setae and with same orientation; 
4 pairs of large, lateroclinate fronto-orbital 
setae; arista plumose, length of branching 
rays nearly equal to twice basal aristal 
width; upturned genal setae lacking, antero- 
clinate genal seta 1, inserted near level of 
anterior margin of eye. 

Thorax: Dorsocentral setae 4 (1 + 3), all 
subequal in size; acrostichal setae small (if 
at all evident), in 2 rows, lacking large pair 
of prescutellar setae; 1 pair of scutellar se- 
tae; anterior supra- alar seta lacking or much 
reduced in length; anterior notopleural seta 
lacking; anepisternal setae pale; katepister- 
nal seta lacking; hindtibia with conspicu- 
ous, rather stout, apical seta anteroventrally; 
apical section of vein M straight. 

Abdomen: Female genital lamellae very 
broad basally, basilateral margins rounded, 
narrowed abruptly near level of cleft, la- 
mellae very narrow from level of cleft to 
apices, with only 1 large stout, acute, ter- 
minal seta at each apex. External male ter- 
minalia: surstylus a single ventral projec- 
tion from ventral margin of epandrium, 
shape like an inverted T or more or less 
triangular, ventrally produced to an anterior 
and posterior angle. 

Remarks. — There are now four Austra- 
lian species of Chaetocanace, including 
three that are newly described. This more 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



than doubles the known species in this ge- 
nus and estabhshes Australia as the center 
for diversity for Chaetocanace. Only C. 
biseta apparently does not occur in Austra- 
ha. 

Key to Australian Species of 
Chaetocanace 

1 . Mesonotum and frons mostly brown 
to blackish brown, subshiny; sursty- 
lus shaped like an inverted T (Fig. 
7), basal portion a gradually nar- 
rowed extension from the ventral 
margin of the epandrium, apical Vs 
projected anteriorly and posteriorly 
C. koongarra, new species 

- Mesonotum and frons brown to 
gray, dull; surstylus shaped other- 
wise 2 

2. Femora mostly yellow; mesonotum 
mostly gray or tannish gray; sursty- 
lus only gradually becoming wider 
ventrally, ventral margin shallowly 
emarginate (Fig. 6) 

C. flavipes, new species 

- At least fore- and hind femora most- 
ly gray; mesonotum mostly grayish 
brown to brown; surstylus becoming 
markedly wider ventrally, more or 
less triangular 3 

3. Anteroventral process of surstylus 
shaped like a long foot (Fig. 8); pos- 
teroventral process distinctly point- 
ed C. longicauda, new species 

- Anteroventral process of surstylus 
not footlike 4 

4. Surstylus with posterodorsal angle 

acutely pointed C. biseta (Hendel) 

Surstylus with posterodorsal angle 
bluntly rounded . . C. brincki Delfinado 



lamadu; figs, of cJ & ? terminalia]. — 
Delfinado & Wirth, 1977:391 [Oriental 
catalog]. — Mathis, 1982:9 [catalog, key]; 
1989a:670 [Australasian/Oceanian cata- 
log]; 1992:5 [world catalog]. 

Australian specimens examined. — New 
South Wales: North Cronulla, 12 Mar 1962, 
D. K. Mc Alpine (1 S; AM). Queensland: 
Cairns (bay shore, puddles), 19-25 Apr 
1957, W. W. Wirth (25 c?, 26 9; AM, 
ANIC, USNM). Eurimbula (mangroves), 
28 Mar 1975, D. K. McAlpine (Id; AM). 
Western Australia: Batten Point (30 km NE 
by E of Borroloola), 18 Apr-30 Oct 1975, 
1976, D. H. Colless, M. S. Upton (4 ?; 
ANIC). 

Distribution. — Australasian/Oceanian: 
Australia (NSW, QLD, WA). Oriental: Phil- 
ippines (Luzon). Sri Lanka. Palearctic: Ja- 
pan (Hokkaido, Honshu, Kyushu, Shikoku), 
Korea (Seoul). 

Diagnosis. — Externally this species is 
very similar to C. biseta (Hendel), and 
many Australian specimens have been mis- 
identified as the latter or as the new species 
described below. I can distinguish between 
C. biseta and this species only on the basis 
of characters of the male terminalia, es- 
pecially the shape of the surstylus (Fig. 5), 
which is more or less triangular in lateral 
view, with the posterodorsal comer broadly 
produced and bluntly rounded. This species 
differs from C. koongarra in having the 
mesonotum lighter in color, mostly dull, 
brown to grayish brown, and only the apical 
tarsomere is brown. 

Remarks. — This species was reported 
previously from Australia (Mathis 1992), 
but only from Queensland. It is now found 
to be more widespread, occurring south- 
ward to New South Wales and westward to 
Western Australia. 



Chaetocanace brincki Delfinado 
Fig. 5 

Chaetocanace brincki Delfinado, 1975:221 
[HT 6 (ZIL); Sri Lanka. Northern Prov- 
ince: Mannar (16 km E), Nay Am at Pal- 



Chaetocanace flavipes, new species 
Fig. 6 

Diagnosis. — This species is distin- 
guished from C. brincki primarily by the 
shape of the external male terminalia, spe- 



VOLUME 109, NUMBER 2 



333 





Figs. 5—6. External male terminalia of Chaetocanace species: 5, epandrium, cercus, and surstylus of C 
brincki, lateral view. 6, epandrium, cercus, and surstylus of C. flavipes, lateral view. Scale = 0. 1 mm. 



cifically the surstylus which in lateral view 
becomes very gradually wider ventrally to 
form short anterior and posterior processes. 

Description. — Small to moderately small 
beach flies, body length 1.8 to 2.3 mm. 

Head: Frons blackish brown and subshi- 
ny on anterior %, portion posterior to an- 
terior ocellus dark brown and less shiny. 
Antenna blackish brown; arista brown. Face 
and gena densely microtomentose, whitish 
to faintly bluish white. 

Thorax: Mesonotum gray, especially lat- 
erally and anteriorly, to mostly grayish 
brown to brown with faint olivaceous col- 
oration, dull; pleural areas generally whitish 
gray; anepisternum faintly bluish; katepi- 
stemum whitish gray. Acrostichal setae 
greatly reduced or lacking, not evident. 
Wing hyaline; costal vein ratio 0.28; M vein 



ratio 0.5. Legs yellow; femora lightly and 
sparsely invested with whitish microtomen- 
tum; apical tarsomere brown. 

Abdomen: External male terminalia as 
follows: surstylus in lateral view (Fig. 6) 
gradually becoming wider ventrally, antero- 
ventral and posteroventral angles produced 
into short processes, posterior process wid- 
er and more bluntly produced; ventral mar- 
gin shallowly emarginate. 

Type material. — The holotype male is la- 
beled "PAPUA NEW GUINEA[,] Central 
Prov[ince]. Lea Lea[,] 6 Oct 1985, J. W 
Ismay (saltpan margin)." Forty paratypes 
(16c?, 249; USNM) bear the same locality 
data as the holotype (dates vary from 23 
Feb to 6 Oct 1985, 1986). Other paratypes 
are as follows: Australia. Western Australia: 
West Kimberley, Cape Bertholet (4 km 



334 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



SSW; at light), 18 Apr 1977, D. H. Colless 
(Id; ANIC). West Kimberley, Cape Ber- 
tholet (8 km S), 18-19 Apr 1977, D. H. 
Colless (1 S, 1 2; ANIC). Drysdale River 
(15.02°S, 126.55°E), 3-8 Aug 1975, I. R. 
B. Common, M. S. Upton (1 S, 2 ?; 
ANIC). The holotype is in excellent con- 
dition, is double mounted (minute nadel in 
block of plastic), and is deposited in the 
USNM. 

Distribution. — This species is known 
thus far from the type locality in Papua 
New Guinea (Central Province) and West- 
ern Australia. 

Etymology. — The specific epithet, flavi- 
pes, refers to the yellow legs, especially the 
femora. 

Chaetocanace koongarra, new species 
Fig. 7 

Diagnosis. — This species is distin- 
guished from C. brincki primarily by the 
shape of the external male terminalia, spe- 
cifically the surstylus which is narrowly 
produced on the basal Vs, gradually tapered 
toward venter, thereafter on apical % rather 
abruptly produced anteriorly and posterior- 
ly to form distinct processes, posterior pro- 
cess shorter and invested with long micro- 
tomentum. 

Description. — Moderately small beach 
flies, body length 2.4 mm. 

Head: Frons blackish brown and subshi- 
ny on anterior 2/3, portion posterior to an- 
terior ocellus dark brown and less shiny. 
Antenna blackish brown; arista brown. Face 
and gena densely microtomentose, whitish 
to faintly bluish white. 

Thorax: Mesonotum blackish brown to 
brown, subshiny, darker and shinier through 
dorsocentral tract of setae, similar to frons; 
anterior supra-alar seta present although 
greatly reduced, less than V2 length of pos- 
terior seta; pleural areas generally whitish 
gray; anepisteraum faintly bluish; katepi- 
sternum whitish gray. Acrostichal setae 
greatly reduced or lacking, not evident. 
Wing hyaline; costal vein ratio 0.22; M vein 



ratio 0.63. Legs mostly yellow; femora 
lightly and sparsely invested with whitish 
to whitish gray microtomentum; apical 1-2 
tarsomeres brown. 

Abdomen: External male terminalia as 
follows: surstylus in lateral view (Fig. 7) 
narrowly produced basally, gradually ta- 
pered toward venter, thereafter rather 
abruptly produced anteriorly and posterior- 
ly to rounded processes; posterior ventral 
process shorter and anterior process, bear- 
ing few setulae but invested with hairlike 
microtomentum; anterior ventral process 
bare. 

Type material. — The holotype male is la- 
beled "[AUSTRALIA.] Koongarra, 15km 
E of Mt. Cahill, N[orthern].T[erritory]. 6- 
9 March 1973[,] D. H. Colless/At light." 
The holotype is in good condition, is double 
mounted (minute nadel in block of poly- 
porus), and is deposited in the ANIC. 

Distribution. — This species is known 
only from the type locality. 

Etymology. — The specific epithet, koon- 
garra, is the name of the type locality and 
is a noun in apposition. 

Chaetocanace longicauda, new species 
Fig. 8 

Diagnosis. — This species is distin- 
guished from C. brincki primarily by the 
shape of the external male terminalia, spe- 
cifically the surstylus which has the antero- 
ventral process greatly produced, elongate, 
forming a footlike process; postero ventral 
process of surstylus distinctly pointed. 

Description. — Moderately small beach 
flies, body length 2.1 to 2.6 mm. 

Head: Frons grayish brown, dull. Anten- 
na blackish brown; arista brown. Face and 
gena densely microtomentose, whitish to 
faintly bluish white. 

Thorax: Mesonotum grayish brown, dull; 
pleural areas generally whitish gray; an- 
epistemum faintly bluish; katepistemum 
whitish gray. Acrostichal setulae in 2 rows; 
anterior supra-alar seta lacking. Wing hya- 
line; costal vein ratio 0.2; M vein ratio 0.5. 



VOLUME 109, NUMBER 2 



335 





Figs. 7-8. External male terminalia of Chaetocanace species: 7, epandrium, cercus, and surstylus of C. 
koonqarra, lateral view. 8, epandrium, cercus, and surstylus of C. longicauda, lateral view. Scale = 0. 1 mm. 



Legs with femora mostly gray, midfemora 
lighter gray; tibiae and tarsi yellow, apical 
1-2 tarsomeres brown. 

Abdomen: External male terminalia as 
follows: surstylus in lateral view (Fig. 8) 
more or less triangular, anteroventral pro- 
cess greatly produced, elongate, shaped like 
a foot, posteroventral angle distinctly point- 
ed, ventral margin deeply sinuous. 

Type material. — The holotype male is la- 
beled "AUSTRALIA: N[orthern]. T[erri- 
tories].; S. Alligator River area, 35kmW 
Jabiru[,] blackhght, 9ivl980 [9 Apr 1980] 
GFHevel & JAFortin." The holotype is in 
fair condition (collected in alcohol, dried, 
then pointed, dorsum appearing "rubbed"), 
is double mounted (glued to a paper trian- 
gle), and is deposited in the ANIC. Fifteen 
paratypes (3 S, 12 9; ANIC, USNM) bear 



the same label data as the holotype. Other 
paratypes are as follows: Australia. Queens- 
land: Cairns, 19 Apr 1957, W. W. Wirth (1 
S; USNM). 

Distribution. — This species is known 
thus far only from Northern Territory (the 
type locality) and Queensland, Australia. 

Etymology. — The specific epithet, longi- 
cauda, is a noun in apposition, referring to 
the long, footlike posteroventral process of 
the surstylus. 

Genus Dynomiella Giordani Soika 

Dynomiella Giordani Soika, 1956:130. 
Type species: Dynomiella arenicola 
Giordani Soika (= Canace stuckenbergi 
Wirth, 1956), by original designation. — 
Mathis, 1982:9-11 [review]; 1992:5 
[world catalog]. 



336 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Canace, in part, of authors. — Wirth, 1956: 
48-51 [revision, key]; 1960:390 [synon- 
ymy of Dynomiella with Canace]. — Co- 
gan, 1980:694 [Afrotropical catalog]. 

Diagnosis. — Resembling Canacea Cres- 
son (Nearctic) but differing from it and oth- 
er genera by the following combination of 
characters. 

Head: Mesofrons similar to parafrons in 
vestiture but densely microtomentose, dull, 
frequently with coloration difference, usu- 
ally mesofrons darker, brown, with 3-5 
large, proclinate setae along lateral margins, 
midportion of mesofrons bare of setae; 
postocellar setae subequal to ocellar setae 
and with similar orientation; 4 large, latero- 
clinate, fronto-orbital setae; arista short, 
generally lacking setulae, apical % to Vi 
bare, stylelike; upturned genal setae 2; an- 
teroclinate genal setae 1. 

Thorax: Dorsocentral setae 4(1 +3), all 
subequal in size; acrostichal setae conspic- 
uous, in 2-4 regular to irregular rows an- 
teriorly, becoming more regular posteriorly, 
middle rows with setulae slightly larger and 
with large pair of prescutellar setae; lateral 
scutellar setae 2 pairs and with several setae 
dorsally; supra-alar seta 1, anterior seta usu- 
ally lacking; 2 notopleural setae; color of 
pleural setulae variable but usually pale, se- 
tae black; postpronotum bare of setulae; 1- 
2 large, anepisternal setae; katepisternal 
seta lacking; forefemur armature variable, 
some species with row of stout, spinelike 
setae anteroventrally; hindtibia lacking api- 
cal seta anteroventrally; apical section of 
vein M straight. 

Abdomen: Female genital lamellae vari- 
able, either only moderately broad basally, 
short, and with lamellar processes over 
three-fourths total length, each process 
gradually tapered to apex; or lamellae very 
broad basally, subtriangular, long, over one- 
third total length, with each lamellar pro- 
cess narrow, parallel sided; in both cases 
apex of each lamellar process bearing 1 
large, stout, moderately acutely to acutely 




Figs. 9—10. External male terminalia of Dynom- 
iella australica: 9, epandrium, cercus, and surstylus, 
lateral view; 10, surstylus, posteroventral view (large 
stilettolike setae along median margin not included). 
Scale = 0.1 mm. 



pointed seta; surstylus with slight, anteri- 
orly curved process, slightly hooklike. 

Dynomiella australica, new species 
Figs. 9-10 

Diagnosis. — This species is similar to D. 
glauca (South African) but differs in having 
only 2 rows of acrostichal setulae; the scu- 
tellum broadly rounded posteriorly, bearing 
a single pair of scutellar setae (the apical 
pair) and sparse setulae on scutellar disc 
that are more concentrated laterally; and 
postpronotum bearing 2 large setae. Fore- 
and midfemora totally lacking prominent 
black setae along the posteroventral and an- 
teroventral surfaces. 

Description. — Moderately small to me- 
dium-sized beach flies, length 2.4 to 3.4 
mm; generally microtomentose, generally 
gray but with some whitish gray to brown- 
ish gray areas, slightly darker dorsally. 

Head: Triangular mesofrons (ocellar tri- 
angle or frontal vittae) well developed, usu- 



VOLUME 109, NUMBER 2 



337 



ally more brown to bronze colored than 
gray to charcoal-gray parafrons, lateral mar- 
gins straight not curved medially, anterior 
angle extended to anterior margin of frons, 
all of median area bare of setae or setulae, 
mesofrontal seta along lateral margins only; 
parafrons bearing sparse, randomly inserted 
setulae, 5-10, none especially well devel- 
oped. Ocelli forming equilateral triangle. 

Thorax: Acrostichal setulae large, in 2 
regular rows, sometimes with 1-2 smaller 
setulae laterad of rows anteriorly; scutellum 
with posterior margin broadly rounded and 
short, not triangular; only apical scutellar 
setae well developed, lacking a basal pair; 
scutellar disc bearing sparse setulae, 5-7, 
these more concentrated laterally; postpron- 
otum bearing 2 large setae, dorsal seta al- 
most as large as posterior seta. Setulae of 
anepisternum and katepisternum fine, pale, 
pilelike, only 2 large setae toward posterior 
margin of anepisternum black, prominent. 
Wing hyaline with faintly brown hues; cos- 
tal vein ratio 0.24; M vein ratio 0.6. Setae 
and setulae of legs pale, mostly white but 
some faintly yellowish white except for 
ventroapical, spinelike black setae of mid- 
tibia; forefemur lacking prominent setae 
along anteroventral surface; midfemur lack- 
ing row of prominent setae along postero- 
ventral surface; tarsi yellow. 

Abdomen: Generally gray; setulae on 
ventral surface of tergites white; tergites 1- 
2 with setulae on dorsum mostly white; 
setulae along posterior margin of 3rd ter- 
gites black; setulae on dorsum, especially 
along posterior margin, of tergites 4-7 and 
epandrium black. External male terminalia 
as follows: epandrium deeply and widely 
cleft posterodorsally; surstylus in lateral 
view (Figs. 8-9) narrow, 3X as long than 
wide, basal % projected directly ventrad 
from ventral margin of epandrium, there- 
after angled anteroventrally, apical Vs much 
narrower, posterior margin along angle 
bearing 4-5 well-developed, stilettolike se- 
tae; anterior margin with moderately long, 
fine setulae; surstylus in posteroventral 
view (Fig. 9) narrow, becoming narrower 



toward apex, median margin mostly 
straight, bearing stilettolike setae, lateral 
margin sinuous, angulate inward at basal Vs, 
thereafter shallowly incurved to apical % 
which is recurved to apex. 

Type material. — The holotype male is la- 
beled "Squeaking Pt. nr Port Sorell, 
Tas[mania, Australia] 24.xi.1968 [24 Nov 
1968] stony beach I. C. Yeo/U.Q.I.C. loan 
724 [pale green on dorsum; number hand- 
written]/HOLOTYPE 6 Dynomiella aus- 
tralica W.N.Mathis [red; gender and species 
name handwritten]." The holotype is dou- 
ble mounted (smaller pin in a long rectan- 
gular block of polyporus), is in good con- 
dition, and is deposited in UQIC. The al- 
lotype female and 23 other paratypes (11 
S, 12 9; UQIC, USNM) bear the same la- 
bel data as the holotype. 

Distribution. — Australasian: Australia 
(TAS). 

Etymology. — The specific epithet, aus- 
tralica, is an adjective and refers to the con- 
tinent where this species occurs. 

Genus Isocanace Mathis 

Isocanace Mathis, 1982:11. Type species: 
Isocanace briani Mathis, by original des- 
ignation. — Mathis, 1992:5-6 [world cat- 
alog]. 

Canace, in part, of authors. — Mathis & 
Wirth, 1979:786. 

Diagnosis. — Resembling Chaetocanace 
Hendel but differing from it and other gen- 
era by the following combination of char- 
acters. 

Head: Mesofrons distinct from parafrons, 
shinier, less microtomentose, with 2-3 
large, lateral, generally proclinate setae; 
postocellar setae smaller than ocellar setae 
and with more proclinate orientation; 4 
pairs of large, lateroclinate, fronto-orbital 
setae; arista plumose, length of branched 
rays varying from approximately subequal 
to nearly twice basal aristal width; upturned 
genal setae 2-3; anteroclinate genal seta 1. 

Thorax: Dorsocentral setae 4(1 +3); ac- 
rostichal setae evident, in 2 rows but lack- 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Figs. 11-18. Scanning electron micrographs of Isocanace albiceps: 11, head, lateral view; 12, gena and 
setae, lateral view; 13, frons, dorsal view; 14, same, left side, dorsal view; 15, ocellar triangle, dorsal view; 16, 
Notopleuron and setae, lateral view; 17, Scutellum, dorsal view; 18, Katepisternum and setae, lateral view. 



ing large pair of prescutellar acrostichal se- 
tae; 2 pairs of scutellar setae and frequently 
with some smaller setae inserted dorsally; 
with only 1 pair of supra-alar setae; 1-2 
notopleural setae, if only 1, anterior seta 
lacking; color of pleural setae variable, pale 
yellow to black; postpronotum bare of setu- 
lae; katepisternal seta present or absent; 1 
large anepisternal seta; hindtibia lacking 
apical seta anteroventrally; apical section of 
vein M straight. 

Abdomen: Female genital lamellae very 
broad basally, basilateral margins rounded, 
narrowed rather abruptly at level of cleft, 
lamellae very narrow from level of cleft to 
apices, with only 1 large, stout, acute ter- 
minal seta at each apex; surstylus quite 



variable, generally slender and with apical 
curvature. 

The albiceps Group 

Diagnosis. — Similar to the briani group 
but differing as follows: Head: mesofrons 
bare in middle; upturned genal setae 2. 
Thorax: katepisternal seta lacking; number 
of acrostichal setae reduced, usually less 
than 10, and usually paired; anterior noto- 
pleural seta subequal to posterior seta. 

Isocanace albiceps (Malloch) 
Figs. 11-18 

Canace albiceps Malloch, 1925:87 [HT 9 
(AM); Austraha. New South Wales: Syd- 
ney]. — Wirth, 1951:262 [review]. 



VOLUME 109, NUMBER 2 



339 





Figs. 19-20. External male terminalia of Isocanace albiceps: 19, epandrium, cercus, and surstylus, lateral 
view; 20, surstylus, lateral view. Scale = 0.1 mm. 



Isocanace albiceps. — Mathis, 1982:18 [ge- 
neric combination]; 1989a:670 [Austral- 
asian/Oceanian catalog]; 1992:6 [world 
catalog]. — Colless & Mc Alpine, 1991: 
779 [fig. of head]. 

Australian specimens examined. — New 
South Wales: Broulee, 17 Sep 1978, Z. Lie- 
pa (2 cJ, 3 9 ; ANIC). Careel Bay, 22 Mar- 
23 Oct 1956, 1962, D. K. McAlpine, W. W. 
Wirth (35 d, 56 9; AM, ANIC, USNM). 
Karuah (inlet, beach), 23 Dec 1968, I. C. 
Yeo (4 d, 9 9 ; UQIC). McCarrs Creek, 20 
Sep 1956, W. W. Wirth (2 $\ USNM). Mer- 
imbula (mangrove flat), 12 Feb 1963, D. K. 
McAlpine (1 9; AM). Mona Vale, 11 Nov 
1956, W W Wirth (1 S; USNM). North 
Cronulla (mangroves), 29 Jan-22 Mar 
1962, D. K. McAlpine (4 cJ, 3 9; AM). 
Putty Beach (near Terrigal), 25 Nov 1987, 
R. Blanche, B. Day, D. K. McAlpine (1 6\ 
AM). Queensland: Deception Bay, 23 May 
1966, Z. Liepa (1 S; ANIC). Tasmania. 
Squeaking Point, near Port Sorell (stony 
beach), 24 Nov 1968, I. C. Yeo (1 d, 8 9; 
UQIC). 

Distribution. — Australasian: Eastern 
Australia (NSW, QLD, TAS). 

Diagnosis. — Specimens of /. albiceps are 
similar to those of the briani group but are 



distinguished by: mesofrons bare in middle; 
postocellar seta short and with more pro- 
clinate orientation; arista with branching 
rays long, some nearly double basal aristal 
width; upcurved genal setae 2; anterior no- 
topleural seta subequal in length to poste- 
rior seta; anepisternal setae pale; katepister- 
num lacking a large seta; surstylus (Figs. 
19-20) comparatively wide in lateral view, 
narrowed subapically, but widened again 
apically and slightly bulbous, with slight 
median projected process, posterior margin 
of surstylus sinuous, anterior margin 
straight. 

Genus Trichocanace Wirth 

Trichocanace Wirth, 1951:252. Type spe- 
cies: Trichocanace sinensis Wirth, by 
original designation and monotypy; 
1964:225-227 [revision; key].— Delfina- 
do & Wirth, 1977:392 [Oriental cata- 
log].— Mathis & Wirth, 1979:795 [diag- 
nosis, discussion]. — Mathis, 1982:20-22 
[review]; 1992:6 [world catalog]. 

Diagnosis. — Resembling Xanthocanace 
Hendel and Chaetocanace Cresson but dif- 
fering from them and other genera by the 
following combination of characters. 



340 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Head: Mesofrons and parafrons dull, 
membranouslike, with fine microtomentose 
vestiture, distinguished from each other by 
color and in one species by density of mi- 
crotomentum, with larger setulae along lat- 
eral margins and a few smaller setulae on 
midportion, but with bare area anterior of 
median ocellus; postocellar setae subequal 
to ocellar setae and with same orientation, 
ocellar setae anterolaterad of ocellar trian- 
gle; 4 pairs of large, lateroclinate, fronto- 
orbital setae; arista with setulae basally, 
setulae not longer than aristal base, apical 
one-fourth or less bare, stylelike; upturned 
genal setae 2, posterior seta directly ventrad 
of midportion of eye, anterior seta aligned 
with anterior margin of eye; anteroclinate 
genal seta present, although small and pale. 

Thorax: Dorsocentral setae 4(1 +3), an- 
terior 2 smaller than posterior setae; acro- 
stichal setulae small, pale, in 4-6 rows, 
lacking large, prescutellar pair of setae; scu- 
tellar setae 1 pair, large, with a few smaller, 
pale setulae dorsally; supra-alar seta 1, an- 
terior seta lacking; anterior-notopleural seta 
lacking; anepisternal and katepisternal setae 
pale, numerous, mostly long and thin, pi- 
lose; katepisternal seta lacking; hindtibia 
without conspicuous, stout, apical seta an- 
teroventrally; apical section of vein M rath- 
er straight, not arcuate. 

Abdomen: Female genital lamellae broad 
basally, basal Va to Vi more or less elliptical, 
with dorsal surface extended, apical Vi to % 
as 2 parallel-sided, narrow processes, each 
bearing 1 large, apical, acutely pointed, 
stout seta and several smaller setae; sursty- 
lus a simple ventral process from epan- 
drium, variously shaped, but usually with 
apex slightly to obviously curved anterior- 

ly- 

Key to Australian Species of 
Trichocanace 

1. Large black species, with heavy 
gray to pearl-gray micro tomentum; 
dorsum of thorax subshiny; frons 
dull black laterally, with broad, me- 



dian, pearl-gray stripe and gray, mi- 
crotomentose, posterior margin; 
both sexes with conspicuous, antero- 
ventral and posteroventral combs, 
each with 6-10 strong, black spine- 
like setae on distal half of forefemur 
T. atra Wirth 

- Dull gray species; frons not black 
striped; forefemur with only antero- 
ventral comb or combs lacking .... 2 

2. Large species, wing length 3.5 mm; 

forefemur lacking combs 

T. sinensis Wirth 

- Small species, wing length 2.4 mm; 
forefemur with inconspicuous, an- 
teroventral comb of 6-8 brownish, 
slender, sharp spinelike setae on dis- 
tal half T. marksae Wirth 



Trichocanace atra Wirth 
Fig. 21 

Trichocanace atra Wirth, 1964:227 [HT S 
(USNM, 67135); Austraha. Queensland: 
Cairns; fig. of S terminalia]. — Delfinado 
& Wirth, 1977:392 [Oriental catalog].— 
Mathis 1982:21 [catalog, key]; 1989a: 
670 [Australasian/Oceanian catalog]; 
1992:6 [world catalog]. 

Australian specimens examined. — North- 
ern Territory: Howard Crossing, 12 May 
1955, L. D. Crawford (1 ?; ANIC). 
Queensland: Bowen (12 mi SE), 6 May 
1955, K. R. Norris, Common (1 9; ANIC, 
allotype). Dunwich, North Stradbroke Is- 
land, 20-21 Apr 1965, G. Monteith (5 ?; 
CNC). Emu Park (NE Rockhampton), 7 
May 1970, Z. Liepa (1 <?; ANIC). Mt. Toz- 
er (11 km ENE, 12°43'S, 143°18'E), 11-16 
Jul 1986, D. H. Colless (1 S\ ANIC). 
Southport, 6 May 1971, B. H. Kay (1 cJ, 1 
$ ; UQIC). Western Australia: Martin's 
Well (1 km S), West Kimberley, 26 Apr 
1977, D. H. Colless (1 c?; ANIC). 

Distribution. — Australasian: Australia 
(NT, QLD, WA). Oriental: Philippines 
(Mindanao), Thailand (Cholburi). 

Diagnosis. — This is the most distinctive 



VOLUME 109, NUMBER 2 



341 




Fig. 21. External male terminalia of Trichocanace 
atra: 21, epandrium and surstylus, lateral view. Scale 
= 0.1 mm. 



species of the genus and is easily distin- 
guished from congeners by the following 
characters: large size (wing length 3.3 mm 
or larger), generally mostly black color with 
dense gray to pearly gray microtomentum; 
dorsum of thorax subshiny; frons dull black 
laterally, with broad, median, pearl-gray 
stripe and gray, microtomentose, posterior 
margin; forefemur of both males and fe- 
males bearing anteroventral and postero- 
ventral comblike rows of 6-10 well-devel- 
oped setae along distal half. 



Trichocanace marksae Wirth 
Figs. 22-31 

Trichocanace marksae Wirth, 1964:226 
[HT S (USNM, 67134); Australia. 
Queensland: Cairns (bayshore); fig. of S 
terminalia]. — Mathis, 1982:21 [catalog, 
key]; 1989a:670 [Australasian/Oceanian 
catalog]; 1992:6 [world catalog]. 

Australian specimens examined. — 
Queensland: Cairns (bay shore), 25 Apr-21 
Dec 1957, 1976, G. F. Hevel, W. W. Wirth 
(6 (J, 22 9; ANIC, USNM). 

Distribution. — Australasian: Eastern 
Austraha (QLD). 

Diagnosis. — This species is the smallest 
of the genus (wing length 2.5 mm or small- 
er) and is otherwise distinguished as fol- 
lows: generally dull gray; frons not striped; 
and forefemur bearing only anteroventral, 
comblike row of 6-8 brownish, slender, 
sharp setae. External male terminalia as fol- 
lows: Epandrium in lateral view (Fig. 31) 
with anterior margin shallowly sinuous, 
posterior margin gently curved, acutely nar- 
rowed dorsally; surstylus in lateral view 
(Fig. 31) narrow, length more than twice 
width, anterior margin more or less straight, 
bearing patch of 4-6 moderately long setu- 
lae near middle, posterior margin rounded, 
ventral margin pointed anteroventrally. 

Trichocanace sinensis Wirth 

Trichocanace sinensis ^ir{h, 1951:253 [HT 
S (BMNH); China. Fukien Province: 
Foochow (= Minhow); fig. of head, 
wing, S terminalia]; 1964:225 [re- 
view].— Delfinado & Wirth, 1977:392 
[Oriental catalog]. — Mathis & Wirth, 
1979:795 [review].— Mathis, 1982:22 
[catalog, key]; 1989a:670 [Australasian/ 
Oceanian catalog]; 1992:6 [world cata- 
log]. 

Australian specimens examined. — 
Queensland: Cairns (bay shore), 19 Apr- 
18-21 Dec 1957, 1976, G. F Hevel, W W 

Wirth {2 6,2 9; ANIC, USNM). Cardwell, 



342 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 




Figs. 22-30. Scanning electron micrographs of Trichocanace marksae: 22, head, lateral view; 23, gena and 
setae, lateral view; 24, frons, dorsal view; 25, same, left side, dorsal view; 26, ocellar triangle, dorsal view; 27, 
antenna, lateral view; 28, scutellum, dorsal view; 29, notopleuron and setae, lateral view; 30, katepisternum and 
setae, lateral view. 



8 Dec 1962, K. R. Norris (9 S, 5 $; 
ANIC). 

Distribution. — Afrotropical: Madagascar 
(sub-Ouest). Australasian: Eastern Australia 
(QLD). Oriental: China (Fukien Province), 
Malaysia (Negri Sembilan), Thailand 
(Bangkok). 

Diagnosis. — This species is large (wing 
length 3.5 mm), similar to T. atra, but dif- 
fers as follows: generally dull gray; frons 
not bearing black stripes; and forefemur 
lacking comblike rows of stout setae. 

Genus Xanthocanace Hendel 

Xanthocanace Hendel, 1914:98. Type spe- 
cies: Canace ranula Loew, by original 
designation. — Malloch, 1924:334 [dis- 
cussion, generic key]. — Cresson, 1936: 
270 [synonymy, discussion]. — Curran, 



1934:357 [generic key].— Wirth, 1951: 
249 [review, key]. — Miyagi, 1963:123 
[review, key]. — Delfinado & Wirth, 
1977:393 [Oriental catalog]. — Mathis, 
1982:22-25 [review]; 1989a:670 [Aus- 
tralasian/Oceanian catalog]; 1992:6-7 
[world catalog]. 

Dinomyia Becker, 1926:107. Type species: 
Canace ranula Loew, by monotypy, pre- 
occupied, Martynov, 1909, and Dyar, 
1919. — Cresson, 1936:270 [synonymy 
with Xanthocanace]. 

Myioblax Enderlein, 1935:235. Type spe- 
cies: Canace ranula Loew, by mono- 
typy. — Cresson, 1936:270 [synonymy 
with Xanthocanace]. 

Diagnosis. — Resembling Trichocanace 
Wirth but differing from it and other genera 
by the following combination of characters. 



VOLUME 109, NUMBER 2 



343 




Fig. 31. External male terminalia of Trichocanace 
marksae: 31, epandrium and surstylus, lateral view. 
Scale = 0.1 mm. 



Head: Mesofrons distinct from parafrons, 
frequently shiny with metallic reflections, 
with numerous, uniformly scattered, pale 
setulae but lacking larger setae along lateral 
margins, anterior margin extended anteri- 
orly beyond level of antennal bases; post- 
ocellar setae subequal to ocellar setae, with 
slightly more divergent orientation; 5-6 
pairs of large to moderately sized, pale, la- 
teroclinate, fronto-orbital setae, anterior 1- 
2 with slight to nearly complete proclinate 
orientation; arista with apical % to Vi bare, 
lacking branching rays, stylelike; upturned 
genal setae lacking, anteroclinate genal se- 
tae 1-3, inserted along anteroventral margin 
of gena. 

Thorax: Dorsocentral setae variable, usu- 



ally only posterior 1-2 setae conspicuously 
larger than surrounding setae, but some 
specimens with up to 6 large setae, some 
presutural, but posterior ones larger; acros- 
tichal setulae in 4 to several rows, these 
more evident anteriorly, lacking large pair 
of prescutellar setae; 2 pairs of scutellar se- 
tae, pale, numerous dorsal setae; 1-2, su- 
pra-alar setae present, anterior one usually 
lacking; anterior notopleural seta present; 
anepisternal setulae pale; katepisternal seta 
lacking; hindtibia without conspicuous, api- 
cal seta anteroventrally; apical section of 
vein M arcuate. 

Abdomen: Female genital lamellae mod- 
erately wide basally, not narrowed abruptly 
near level of cleft, bearing only 1 large, 
stout, acute terminal seta at each apex; sur- 
stylus a simple ventral projection from 
epandrium, tapered gradually, apex acute to 
blunt. 

Key to Australian species of 
Xanthocanace 

1. Mesofrons of male thinly microto- 
mentose, somewhat dull, but with 
some subshiny, metallic luster; 
mesofrons of female entirely silvery 

gray microtomentose, dull 

X. collessi, new species 

- Mesofrons of male mostly shiny, 
very thinly microtomentose; meso- 
frons of female mostly dull, micro- 
tomentose, with some brownish col- 
oration medially, lateral margins 
gray X. nigrifrons Malloch 



Xanthocanace nigrifrons Malloch 
Figs. 32-40 

Xanthocanace nigrifrons Malloch, 1924: 
334 [HT S (AM); Australia. New South 
Wales: Woy Woy; figs, of head]. — Wirth, 
1951:250 [review].— Miyagi, 1963:125- 
126 [distribution, key].— Griffiths, 1972: 
256 [discussion of S terminalia]. — Math- 
is, 1982:24 [catalog, key]; 1989a:670 



344 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Figs. 32-39. Scanning electron micrographs of Xanthocanace nigrifrons: 32, head, lateral view; 33, gena 
and setae, lateral view; 34, frons, dorsal view; 35, Same, left side, dorsal view; 36, ocellar triangle, dorsal view; 
37, antenna, lateral view; 38, notopleuron and setae, lateral view; 39, katepisternum and setae, lateral view. 



[Australasian/Oceanian catalog]; 1992:7 
[world catalog]. 

Australian specimens examined. — New 
South Wales: Angourie, 26 Jan 1980, B. J. 
Day, D. K. McAlpine (2 2; AM). Narra- 
been Lagoon (tidal flat), 12 Oct-5 Dec 
1956, W. W. Wirth (48 c?, 49 $; ANIC, 
USNM). Queensland: MacKay Harbour (on 
sand of beach, high tide mark), MacKay, 25 
Jul 1987, R. Jones (1 6; UQIC). Miriam 
Vale District, Eurimbula (sandy pool on 
beach), 28 Mar 1975, D. K. McAlpine (4 
S, 12 2; AM, USNM). Tasmania: Squeak- 
ing Point near Port Sorell (stony beach), 24 
Nov 1968, I. C. Yeo (18 c?, 16 ?; UQIC). 
Western Australia: Cape Bertholet (4 km 
SSE), West Kimberley, 19 Apr 1977, D. H. 
Colless (IS; ANIC). 



Distribution. — Australasian: Australia 
(NSW, QLD, TAS, WA). 

Diagnosis. — This species is distinguished 
from congeners by the following combina- 
tion of characters: length over 2.5 mm; me- 
sofrons of male very thinly microtomentose, 
subshiny, bluish black; mesofrons of female 
almost entirely microtomentose, dull; mid- 
femur of male lacking posteroventral comb 
of 5-10 small, black setae on distal half; 
femora with at least apical half yellow. Ex- 
ternal male terminalia as follows: surstylus 
in lateral view (Fig. 40) bearing short and 
moderately long setulae, shape narrow, twice 
as long as wide, basal % a gradually narrow- 
ing straight process from ventral epandrial 
margin, thereafter curved abruptly poster- 
oventrally as a much narrower, digitiform 
process, apex bluntly rounded. 



VOLUME 109, NUMBER 2 



345 





Figs. 40-42. External male terminalia of Xanthocanace species: 40, surstylus of X. nigrifrons, lateral view. 
41, epandrium, cercus, and surstylus of X. collessi, lateral view; 42, surstylus of X. collessi, posterior view (setae 
and setulae not included). Scale = 0.1 mm. 



Remarks. — I examined a single female as 
part of this study from Victoria (Mallacoota 
Inlet, SE Genoa, 4 Aug 1973, Z. Liepa 
(ANIC)) that is very similar to and may be 
conspecific with this species. This female is 
darker in coloration, particularly on the 
mesonotum, and the mesofrons also has 
darker hues with slightly more metallic lus- 
ter shining through the micro tomentum. 
Additional specimens, especially males, 
will be needed to resolve the status of this 
population. 

Xanthocanace collessi, new species 
Figs. 41-42 

Diagnosis. — This species is similar to X. 
sabroskyi Mathis & Freidberg and especial- 
ly X. zeylanica Delfinado and is distin- 
guished from either and other congeners by 
the following combination of characters: 
Mesofrons of male thinly microtomentose, 
with some subshiny, metallic luster; meso- 
frons of female entirely microtomentose sil- 
very gray, dull; femora mostly whitish gray 
except for yellow apices; tibiae yellow. 



Description. — Moderately small to me- 
dium-sized beach flies, length 2.2 to 3.3 
mm: body mostly silvery white to slightly 
metallic bluish to whitish gray, darker dor- 
sally with some subshiny areas with metal- 
lic luster. 

Head: Mesofrons of male thinly micro- 
tomentose, somewhat dull, partially sub- 
shiny with some metallic gray to silvery 
gray luster; mesofrons of female entirely 
silvery gray microtomentose, dull; meso- 
frontal setulae abundant, conspicuous. Face, 
gena, and clypeus concolorous, silvery 
white, densely microtomentose. 

Thorax: Mesonotum silvery gray, fre- 
quently with some faint bluish coloration, 
slightly subshiny, with weak metallic luster; 
pleural areas lighter, becoming almost com- 
pletely white. Wing faintly milky white; 
costal vein ratio 0.28; M vein ratio 0.52. 
Legs: midfemur of male lacking row of 5- 
10 closely set black setae along apical half 
of posteroventral surface; femora mostly 
whitish gray to gray, except for yellow api- 
ces; tibiae yellow to whitish yellow. 



346 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Abdomen: Dorsum concolorous with 
mesonotum. Male preabdomen as follows: 
sternite 3 generally rectangular with round- 
ed corners, slightly narrower anteriorly than 
posteriorly; sternite 4 nearly as wide as 
long, anterior margin narrower than poste- 
rior margin, posterior margin shallowly and 
narrowly notched medially; sternite 5 
broadly V-shaped with arms angled poste- 
riorly, posterior margin fused with more 
lightly sclerotized 6th sternite. External 
male terminalia as follows: epandrium with 
a deep, narrow, posteromedian cleft dorsal- 
ly, not bearing long black setae with pos- 
terior orientation; surstylus in lateral view 
(Fig. 41) nearly as wide as long, anteroven- 
tral margin rounded, posteroventral margin 
produced ventrally to a pointed projection, 
posterior margin shallowly sinuous; sursty- 
lus in posterior view (Fig. 42) very narrow, 
apical % as a slender slipper, with apex ori- 
ented posteromedially, ventral margin bear- 
ing fringe of short setulae. 

Type material. — The holotype male is la- 
beled "[AUSTRALIA] 5 km SSW of Cape 
Bertholet WA West Kimberley 21 Apr 1977 
D. H. Colless/At light/HOLOTYPE S Xan- 
thocanace collessi W. N. Mathis [red; gen- 
der and species name handwritten]." The 
holotype is double mounted (smaller pin in 
a block of polyporus), is in good condition, 
and is deposited in ANIC. The female al- 
lotype and three other paratypes ( 9 ; ANIC) 
bear the same label data as the holotype. 
Other paratypes are as follows: Western 
Australia: Cape Bertholet (8 km S), West 
Kimberley, 17 Apr 1977, D. H. Colless (2 
(?, 28 $; ANIC, USNM). Cape Bertholet 
(3 km S), West Kimberley, 20 Apr 1977, 
D. H. Colless (2 9 ; ANIC). Cape Bertholet 
(4 km SSW), West Kimberley, 18 Apr 
1977, D. H. Colless (2 9 ; ANIC). 

Distribution. — Australasian: Australia 
(WA). 

Etymology. — The specific epithet, colles- 
si, is a genitive patronym to recognize the 
many contributions of Donald H. Colless to 
the study of Diptera, the Australian fauna 



in particular. He also collected all speci- 
mens of the type series. 

New Genus 

Diagnosis. — Resembling Trichocanace 
Hendel and Chaetocanace Cresson but dif- 
fering from them and other genera by the 
following combination of characters. 

Head: Mesofrons and parafrons dull, 
membranelike, with dense microtomentose 
vestiture, distinguished from each other by 
color and density of microtomentum, with 
larger setulae along lateral margins and a 
few smaller setulae on midportion, but with 
bare area anterior of median ocellus; post- 
ocellar setae subequal to ocellar setae and 
with same orientation, ocellar setae inserted 
anterolaterad of ocellar triangle; 3 large, la- 
teroclinate, fronto-orbital setae; arista with 
basal segment bare, apical segments bearing 
tiny hairs; genal setae 3, anteroclinate and 
slightly upturned apically, aligned horizon- 
tally. 

Thorax: Dorsocentral setae 6 (2 + 4), an- 
terior seta smaller than posterior setae, 5th 
seta inserted slightly mediad to alignment 
of anterior setae, posterior seta inserted 
slightly lateral of alignment; acrostichal 
setulae lacking anteriorly but with a prescu- 
tellar pair of setae; lateral scutellar setae 2; 
scutellar disc with a few setulae, 3-5; su- 
pra-alar setae 1 large and 1-2 smaller setae 
anteriad; notopleural setae 2, both well de- 
veloped; anepi sternum with 2 larger setae 
along posterior margin, 2-3 smaller setae 
below and a few setulae medially; katepi- 
sternal seta lacking. Apical section of vein 
M rather straight, very shallowly sinuous, 
not arcuate. Hindtibia without a conspicu- 
ous, stout, apical seta anteroventrally. 

Abdomen: Female genital lamellae re- 
duced and lacking prominent setae apically 
and subapically. 

New Species of New Genus 

Australian specimens examined. — South 
Australia: Coward Springs (5 km NW), 



VOLUME 109, NUMBER 2 



347 



near Lake Eyre, 22 Sep 1972, Z. Liepa (1 
9 ; ANIC). 

Acknowledgments 

For the loan of specimens, I thank the fol- 
lowing individuals and their sponsoring in- 
stitutions: Margaret A. Schneider (UQIC), 
David K. Mc Alpine (AM), and Peter S. 
Cranston (ANIC). Susann Braden assisted 
with the preparation of the scanning electron 
micrographs, and Victor Krantz assisted 
with production of the photographs. The pen 
and ink illustrations were inked by Elaine R. 
S. Hodges. For reviewing a draft of this pa- 
per, I thank Curtis W. Sabrosky, Norman E. 
Woodley, and Oliver S. Flint, Jr. 

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Miyagi, I. 1963. Notes on Korean species of the Can- 

aceidae, with descriptions of two new species 

(Diptera: Canaceidae). — Insecta Matsumurana 

26(2): 122-126. 
Wirth, WW 1951. A revision of the dipterous family 

Canaceidae.-Occasional Papers of the B. P. 

Bishop Museum 20(14):245-275. 



. 1956. New Species and Records of South 

African Canaceidae (Diptera). — Journal of the 
Entomological Society of South Africa 19:47- 
51. 

. 1960. Chapter XIX. Diptera (Brachycera): 

Canaceidae and Ephydridae. In B. Hanstrom, P. 
Brinck, and G. Rudebeck, eds.. South African 
Animal Life Results of the Lund University Ex- 
pedition in 1950-1951. Vol. 7:390-396. Lun- 
dae. 

. 1964. New species and records of the genus 

Trichocanace Wirth (Diptera, Canaceidae). — 
Pacific Insects 6:225-227. 

. 1987. Canacidae. 102 [chapter]. Pp. 1079- 

1083 in J. F. McAlpine, ed.. Manual of Nearctic 
Diptera. Vol. 2. Ottawa: Research Branch, Ag- 
riculture Canada. Monograph No. 28, 1332 pp. 

Womersley, H. 1937. Diptera, British, Australian, and 
New Zealand Antarctic Research Expedition 
Reports, series B, 4(3):59-79. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):349-352. 1996. 

Marine fossil shark (Chondrichthyes) from nonmarine Eocene 
sediments, northeastern Kazakhstan 

Spencer G. Lucas, Robert J. Emry, and Robert W. Purdy 

(SGL) New Mexico Museum of History and Science, 

1801 Mountain Road N.W., Albuquerque, New Mexico, 87104, U.S.A. 

(RJE, RWP) Department of Paleobiology, National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — A single tooth of the sand tiger shark, Carcharias sp., was col- 
lected from the upper Eocene Kusto svita in the Zaysan basin of northeastern 
Kazakhstan. The Kusto svita is nonmarine lacustrine strata with an extensive 
nonmarine biota of plants (especially charophytes) and terrestrial vertebrates 
(especially mammals). The shark tooth was apparently carried to the site by a 
predator/scavenger from the then-nearest seaway, more than 500 km to the 
northwest. 



The Zaysan basin of northeastern Ka- 
zakhstan (Fig. 1) has an Upper Cretaceous- 
Miocene sedimentary fill dominated by 
freshwater shales, siltstones and fine sand- 
stones deposited in and around ancient 
Lake Zaysan (Borisov 1963, Verzilin et al. 
1980). The middle Eocene-middle Miocene 
strata here produce rich fossil assemblages 
of freshwater and terrestrial plants (charo- 
phytes and angiosperm leaves), inverte- 
brates (ostracods, gastropods, unionid bi- 
valves) and vertebrates (amiid and teleost 
fishes, salamanders, turtles, crocodylians, 
birds and mammals). We were thus sur- 
prised to find a single tooth of a marine 
selachian at a late Eocene site in the Zaysan 
basin. Here we document this fossil and 
suggest a probable mechanism for its trans- 
port from the marine to nonmarine environ- 
ment. 



the eastern bank of the Sarybulak River, 
downstream from the well known Eocene 
mammal locality called Sunduk (Russell & 
Zhai 1987, fig. 84). Locality K12 is in the 
Kusto svita in a bed of silty and sandy mud- 
stone that is pale olive with dark yellowish 
orange limonitic mottling. This bed is 8 m 
below the top of the Kusto svita, which is 
approximately 50 m thick in this area. 

Locality K12 produced an extensive ver- 
tebrate fossil assemblage now under study 
that includes an amiid fish, a trionychid 
turtle, a crocodylian, cylindrodontid and 
cricetid rodents, the hyaenodontid creodont 
Hyaenodon, and the anthracothere Elome- 
ryx. Adjacent strata produce numerous 
charophytes (Kyansep-Romashkina 1980). 
Fossil mammals indicate the Kusto svita is 
of late Eocene age (Ergilian land-mammal 
"age" of Russell & Zhai 1987). 



Occurrence 

During the summer of 1993, the fossil 
selachian tooth, KAN (Institute of Zoology, 
Kazakh Academy of Sciences, Almaty) 35 
(12) 1660, was collected from screen wash- 
ed sediment at our locality K12 (UTM 
387305E, 5260046N, zone 45), which is on 



Description and Identification 

KAN 35 (12) 1660 (Fig. 2). A single- 
cusped tooth missing much of root, which 
is heavily abraded. Crown tall (total length 
at least 21.9 mm), narrow, sharply pointed 
blade. Crown with distinctly sigmoidal out- 
line in anterior view. Two cutting edges of 



350 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Miles 


100 


200 




300 


Kilometers 


100 200 


300 


400 


500 



Russia 




r'l 



Novosibirsk 



-^etropavlovsk' 'm . ^ ,y^^ 




.j"c".XC^-l China 

V. Tajikistan ^. 
f Afganistan ^' Pakistan 



Fig. L Map of Kazakhstan showing location of Zaysan basin in the northeastern part of the country. 



crown forming sharp, nearly parallel cari- 
nae extending almost to root. No lateral 
cusplets preserved, but crown base is miss- 
ing where these would have been, if present 
at all. Crown enamel lingually smooth to 
very weakly striated longitudinally. Impos- 
sible to determine if root had transverse 
groove. Lingual face of root smooth and 
convex. Root not massive, but full extent 
cannot be determined. 

This tooth is identical in form to the sec- 
ond upper anterior teeth of several kinds of 
lamniform sharks. Of the lamniform sharks 



with elongate anterior teeth, only the spe- 
cies of Odontaspis, Carcharias and Isurus 
have second upper anterior teeth that re- 
semble KAN 35 (12) 1660. Unlike KAN 35 
(12) 1660, the second upper anterior teeth 
of Odontaspis have cutting edges extending 
only down about one-half to two-thirds the 
distance from the apex of the crown. How- 
ever, the lack of the entire root, so that the 
presence of lateral cusplets and a transverse 
groove cannot be established, make it pos- 
sible that this tooth could be that of a mako, 
Isurus, though the presence of faint stria- 



VOLUME 109, NUMBER 2 



351 





Fig. 2. Photograph of KAN 35 (12) 1660, incomplete tooth of Carcharias sp. from the Kusto svita in the 
Zaysan basin of northeastern Kazakhstan; A, labial; B, lingual; C, anterior. Approximately X2; scale in mm. 



tions on the lingual surface of the crown is 
characteristic of Carcharias, not Isurus. 
Closest resemblance of KAN 35 (12) 1660 
is to anterior teeth of the sand tiger shark 
Carcharias (see Applegate 1965), to which 
we assign it as Carcharias sp. 

Discussion 

The species of Carcharias are marine 
sharks with a temporal range extending 
back to the beginning of the Late Creta- 
ceous (Cenomanian) (Cappetta 1987, Wel- 
ton & Parish 1993). The freshwater origin 
and fossil biota of the Kusto svita at site 
K12 excludes the possibility of the Car- 
charias tooth representing an animal that 
lived at the site, so it is not autochthonous 
to the deposit. Its allochthonous origin 
could not be as a fossil reworked from older 
sediments, simply because no such marine 
strata are present in or around the Zaysan 
basin (Borisov 1963). The youngest marine 
strata in this part of Kazakhstan are of 
Permian age, much older than the first 
known occurrence of Carcharias. 

It seems most likely that this allochtho- 
nous shark tooth was brought to the site by 
a biological agent. During the late Eocene, 
the nearest marine water was the western 
Siberian seaway, with its eastern shoreline 



to the northwest of Semipalitinsk, about 
500 km to the northwest of locality K12 
(Tsekhovsky 1987, fig. 42). Sand tiger 
sharks are nearshore marine sharks that 
would have inhabited the shallow waters 
along the margin of such a seaway. Indeed, 
fossils of Carcharias and other lamniform 
sharks are known from Eocene-Oligocene 
strata in central and western Kazakhstan 
(GHkman 1964). 

We propose that from this seaway a pred- 
ator/scavenger — a crocodyle, bird or mam- 
mal — somehow carried the tooth to the site 
(probably by consumption), where it be- 
came fossilized out of context. At present 
no other explanation fits the data. The iso- 
lated nature of the tooth, the close associ- 
ation of this marine shark fossil with fresh- 
water fossils and the lack of a sedimentary 
source from which it could have been re- 
worked, make transport by a biological 
agent, probably predator/scavenger, the 
only reasonable explanation. To our knowl- 
edge, this is the first case in the fossil record 
of a probable predator/scavenger transport 
of a shark tooth from a marine environment 
to a nonmarine setting. 

Acknowledgments 

The National Geographic Society (Grant 
5412-95) and the Charles D. Walcott Fund 



352 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



of the Smithsonian Institution supported 
this research. Lyuba Tyutkova found local- 
ity K12, and several Kazakh, Georgian and 
American colleagues assisted in the field. V. 
Springer and J. Tyler provided helpful re- 
views of the manuscript. 

Literature Cited 

Applegate, S. P. 1965. Tooth terminology and varia- 
tion in sharks with special reference to the sand 
shark, Carcharias tauriis Rafinesque. — Los An- 
geles County Museum Contributions in Science 
86:1-18. 

Borisov, B. A. 1963. The stratigraphy of the Upper 
Cretaceous and of the Paleogene-Neogene of 
the Zayzan depression [in Russian]. — Trudy 
VSEGEI i Gosudarstveny Geologicheskiy 
Komineta, new series 94:11-75. 

Cappetta, H. 1987. Chondrichthyes II, Mesozoic and 
Cenozoic Elasmobranchii. — Handbook of Pa- 
leoichthyology 3B: 1-193. 

Glikman, L. S. 1964. Paleogene sharks and their 
stratigraphic significance [in Russian]. — Isda- 
telstvo Nauk, Leningrad, 228 pp. 



Kyansep-Romashkina, N. P. 1980. Charophytic algae. 
Pp. 97-124 in G. G. Martinson and N. P Kyan- 
sep-Romashkina, eds., Paleolimnology of Zay- 
san [in Russian]. Isdatelstvo Nauk, Leningrad. 

Russell, D. E., & R. Zhai. 1987. The paleogene of 
Asia: mammals and stratigraphy. — Memoires 
du Museum National d' Historic Naturelle, Sci- 
ences de la Terre 52:1-487. 

Tsekhovsky, Yu. G. 1987. Sedimento- and lithoge- 
nesis of humid red rocks at the transition from 
Cretaceous to Paleogene in Kazakhstan [in Rus- 
sian]. — Akademiya Nauk SSSR Ordena Trudo- 
vo Krasnovo Znameni Geologicheskii Institut 
Trudy 423:1-190. 

Verzilin, N. N., N. P. Kyansep-Romashkina, O. A. Mi- 
ronenko, E. C. Stankevich, N. V. Tolstikova, V. 
M. Chkhikvadze, & N. Shevyreva. 1980. His- 
tory of the development of the lake basin during 
Late Cretaceous and Paleogene time. Pp. 125- 
145 in G. G. Martinson and N. P. Kyansep-Ro- 
mashkina, eds., Paleolimnology of Zaysan [in 
Russian]. Isdatelstvo Nauk, Leningrad. 

Welton, B. J., & R. E Farish. 1993. The collector's 
guide to fossil sharks and rays from the Creta- 
ceous of Texas. Before Time, Lewisville, Texas, 
204 pp. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):353-365. 1996. 

Pholidichthys unguis, a new species of pholidichthyid fish from 
Northern Territory and Western AustraUa 

Victor G. Springer and Helen K. Larson 

(VGS) Division of Fishes, National Museum of Natural History, 

Washington, D.C. 20560, U.S.A; 

(HKL) Museum and Art Gallery of the Northern Territory, 

GPO Box 4646, Darwin NT 0801, Australia 

Abstract. — Pholidichthys anguis is only the second species known in the 
family Pholidichthyidae. It differs from P. leucotaenia in having 87-98 dorsal- 
fin rays (vs. 66-79), 70-81 anal-fin rays (vs. 49-62), 90-101 total vertebrae 
(vs. 71-79), a more slender body, and a shorter head and maxillary. In addition, 
the color pattern of juveniles (—50-103 mm SL) differ in having the depth of 
the slender dark stripe on midside at vertical from anal-fin origin about 1 mm, 
usually much less (vs. depth 2.5-3.3 mm). Study of whole specimens and 
skeletal preparations of P. anguis indicates that the species exhibits the same 
familial specializations as P. leucotaenia. The two species are distributed al- 
lopatrically. We hypothesize that the common ancestor of the two species of 
Pholidichthys had a Tethyan distribution exclusive of Australia-New Guinea 
(ANG). When ANG collided with SE Asia, the ancestor invaded ANG. During 
a subsequent interglacial period, ecological factors contributed to the isolation 
of the Australian portion of the ancestral population from the New Guinea 
portion, and permitted divergence of the isolates. 



Pholidichthys and its only included spe- 
cies, P. leucotaenia, sole representative of 
the Pholidichthyidae, were described by 
Bleeker (1856) from Boeroe (= Buru), In- 
donesia. A purported second species, P. an- 
guilliformis, was described by Lockington 
(1882) from the Gulf of California. The ho- 
lotype of P. anguilliformis has not been 
seen since its description, but characters 
given for the species (especially its all spi- 
nous dorsal fin) and its type locality, prob- 
ably indicate that it is not congeneric or 
confamilial with Pholidichthys Bleeker. 
Larson {in Trnski et al. 1989) indicated that 
an undescribed species of Pholidichthys 
(whose description is the main purpose of 
the present paper) also exists in the Indo- 
Pacific. Her report was based on specimens 
brought to her attention by prawn research- 
ers of the Northern Territory Fisheries Di- 
vision, Darwin. We know of no other de- 



scribed or undescribed taxa that are refer- 
able to the Pholidichthyidae. 

In the last major study of Pholidichthys, 
Springer & Freihofer (1976) reviewed the 
literature on P. leucotaenia and described 
aspects of its osteology, neurology, and on- 
togenetic color-pattern changes. They also 
remarked on its behavior in aquaria, plotted 
its geographic distribution, and discussed 
its possible familial interrelationships. 

Materials and Methods 

Specimens of the new species are listed 
in the description. All but one specimen, an 
adult female, 245 mm SL, which we des- 
ignate holotype, are small juveniles (most 
less than 70 mm SL, none between 103 mm 
and 245 mm) obtained from trawl hauls. 
Most of the specimens are curled and twist- 
ed and do not permit accurate SL measure- 



354 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



ments to be made. We arbitrarily selected a 
few specimens in good condition on which 
to take measurements (Table 1, Fig. 6). 
Nevertheless a cursory examination of 
many specimens convinces us that the main 
proportional differences we stress will hold 
when more specimens in good condition 
become available for measurement. 

Comparative material of Pholidichthys 
leucotaenia is that listed by Springer & 
Freihofer (1976:2) and more recent material 
or records as follows: New Caledonia (Bur- 
gess & Axelrod 1975: fig. 65, photograph; 
see discussion in Springer 1982:71); Solo- 
mon Islands: Marau Island (Burgess & Ax- 
elrod (1975:fig. 64, photograph). Indonesia: 
Flores (Kuiter 1992:4, fig. E, and J. E. 
Randall, pers. comm.), Saparua (USNM 
210334), Tulangbesi Islands (J. E. Randall, 
photograph, pers, comm.); Philippines: 
Tawi Tawi (USNM 122340, 150828, 
151178), Jolo (USNM 122339), Apo 
(USNM 289924), Luzon (questionable rec- 
ord based on aquarium dealer reports). Pa- 
pua New Guinea: Madang and vicinity 
(AMS 1.34712.007, USNM 258321), 
d'Entrecasteaux Islands (J. E. Randall, pho- 
tograph, pers. comm.). Singapore (ZMUC 
P.75457, obtained from an aquarium deal- 
er). 

Dorsal-, anal-, and caudal-fin rays, ver- 
tebrae, pleural, and epineural counts were 
made from radiographs (and from the 3 
cleared-and-stained specimens). Many 
more specimens were radiographed than is 
indicated by the numbers of specimens re- 
ported in Table 2, but because of poor os- 
sification or fineness of structure, we were 
unable to obtain all counts for all speci- 
mens, and for some we were unable to 
make any counts. Specimens were cleared 
with trypsin and counterstained with aliza- 
rin and alcian blue. Dorsal- and anal-fin ray 
counts include all elements (last two rays 
counted as two — last ray not split to base). 
Precaudal vertebrae are those anterior ver- 
tebrae lacking a hemal spine. Measurements 
were made with dial calipers and recorded 
to three significant figures. Standard length 



(SL) was taken from the midtip of the upper 
lip to the midbase of the caudal fin; head 
length (HL), from the midtip of the upper 
lip to the posteriormost edge of the oper- 
culum; snout length, from the midtip of the 
upper lip to the anteriormost edge of the 
orbit; upper jaw length, from the midtip of 
the upper lip to the posteriormost edge of 
the maxilla; predorsal length, from the mid- 
tip of the upper lip to the anterior base of 
the first dorsal-fin ray; orbital diameter is 
the greatest horizontal diameter; postorbital 
head length was taken from the posterior- 
most edge of the orbit to the posteriormost 
edge of the operculum; body depth was 
measured vertically at the anal-fin origin; 
mid-lateral dark stripe depth was measured 
where the stripe crossed a vertical from the 
anus; caudal-fin length is the length of the 
longest caudal-fin ray. Regression formulae 
and plotted curves are the products of a 
computer software program, PSI-Plot, ver- 
sion 3 (Poly Software International, P.O. 
Box 526368, Salt Lake City, UT 84152). 
Institutional abbreviations are those given 
by Leviton et al. (1985). 

Pholidichthys unguis, new species 
Figs. 1-3 

Holotype.—NTM S. 11799-001, mature 
female, 245 mm TL, WSW of Angurugu, 
Groote Eylandt, Northern Territory, Austra- 
lia (14°05'S, 136°15'E), 19 m, 30 June 
1984, Northern Territory Fisheries. 

Paratypes. — 130 specimens (—50-103 
mm SL): AMS 1.36375-001 (3), ANSP 
173800 (3), BMNH 1995.8.14.1-3 (3), 
BPBM 36786 (3), CAS 82409 (3), MNHN 
1995-0896-0898 (3), NTM S. 13530-003 
(26), QM 1.30118 (3), ROM 69314 (3), 
USNM 337859 (9) and WAM R3 1012-001 
(3), all with same data: Arafura Sea, North- 
ern Territory, North Goulburn Island 
(10°23'49"S, 135°43'53"E), depth 53 m, 23 
Sept 1992. NTM S. 13529-001 (49) and 
USNM 337860 (15, including 3 cleared and 
stained), Arafura Sea, Northern Territory, 
W of Cape Wessel (10°57.2'S, 136°06.2'E), 



VOLUME 109, NUMBER 2 



355 



depth 42-43 m, 22 Sept 1992. NTM S- 
13039-001 (3), Arafura Sea, Northern Ter- 
ritory, N of Melville Island (10°38'S, 
130°52'E), 17 June 1989. NTM S-13339- 
002 (1), Timor Sea, Western Australia, Jo- 
seph Bonaparte Gulf (13°07'S, 128°56'E), 
depth 70 m, 23 Nov 1990. 

Diagnosis. — A species of Pholidichthys 
with: 87-98 dorsal-fin rays; 70-81 anal-fin 
rays; 90-101 total vertebrae; depth of slen- 
der, dark stripe on midside of juveniles (up 
to at least 103 mm SL) 1.0 mm or less at 
vertical from anal-fin origin. 

Description (Tables 1-2). — Dorsal fin 
87-98, all rays segmented, all but a few of 
the most anterior and posterior rays 
branched (unable to determine condition in 
holotype), anteriormost ray not supernu- 
merary, last ray not split to base, stay 
(Johnson 1984:caption of table 120) pres- 
ent. Supraneurals or predorsal bones absent. 

Anal fin 70-81 (Table 2), all rays seg- 
mented, all but a few of the most anterior 
and posterior rays branched (unable to de- 
termine condition in holotype), anterior- 
most ray apparently supernumerary (ante- 
riormost pterygiophore appears to be fusion 
of two pterygiophores), last ray not split to 
base, stay present; 7-10, usually 8 or 9, 
pterygiophores anterior to first hemal spine. 

Vertebrae 23-28 + 64-75 = 90-101 (Ta- 
ble 2); posteriormost pleural rib articulating 
with next- to posteriormost or posteriormost 
precaudal centrum; epineurals 18-22. 

Pectoral-fin rays 14 or 15 (14 in 16 of 17 
specimens checked), dorsal- and ventral- 
most two rays simple, others branched. 

Pelvic-fin small, rays 1,2 or 1,3; spine 
greatly reduced, visible only in osteological 
preparations; third (innermost) segmented 
ray greatly reduced when present, usually 
visible only in osteological preparations; 
segmented rays all simple; basipterygia 
abutting posteriorly, widely separated an- 
teriorly; rays supported by an almost spher- 
ical cartilaginous process at ventroposterior 
end of basipterygium. 

Caudal-fin rays 10, all simple, 5 rays 
above and 5 below diastema, parhypural 



and hypurals fused into single plate, no pro- 
current rays, no epurals. 

Gillrakers on first arch (epibranchial-an- 
gle-hypobranchial; left side/right side): 5-1- 
8/5-1-8 (holotype); 5-1-9/5-1-8, 5-1-9/5-1- 
9, 6-1-9/6-1-10 (three cleared-and-stained 
specimens, left & right sides). 

One nostril (anterior) each side; number 
and distribution of sensory pores as in P. 
leucotaenia (Springer & Freihofer 1976: 
figs. 12 and 14), except that interorbital 
commissural pore (Springer & Freihofer 
1976:fig. 12) varying from present and ob- 
vious to absent. 

Teeth: In general, all teeth are moderately 
laterally compressed canines with slightly 
recurved tips. In juveniles, —80 mm SL, 
upper teeth on each side of each jaw ar- 
ranged in 3 linear rows; most posterior row 
with smallest teeth, consisting of about 10 
teeth, of which anteriormost 2 are over- 
lapped externally by posteriormost 1 or 2 
teeth of next row anterior, which consists of 
about 8 slightly larger teeth, of which an- 
teriormost 2 or 3 are overlapped externally 
by posteriormost 2 or 3 teeth of next row 
anterior, which consists of about 6 teeth, 
which are initially larger but decrease in 
size and become misaligned anteriorly; ex- 
ternal to small, misaligned teeth of previous 
row are largest 4 teeth. Anterior, largest 
teeth of upper jaw larger than those of low- 
er jaw. In the holotype, the tooth rows have 
been disrupted and there appears to be 1 
row of teeth posteriorly grading into 3 ir- 
regular rows anteriorly, with the outermost 
anterior teeth the largest. 

Measurements of certain body characters 
are given in Table 1 , and four of these char- 
acters are plotted (Fig. 6) against SL with 
their best fitted regression curve (usually a 
power curve) for comparison with the same 
characters in P. leucotaenia. 

Color pattern (Figs. 1-3): All but 1 of the 
available specimens are juveniles of about 
25 to 103 mm SL. The color pattern of the 
juveniles is fairly consistent, but the dark 
markings we describe vary from faintly to 
darkly dusky, and may be interrupted. We 



356 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 1 . — Measurements in millimeters (and as percent SL) for certain characters of the species of Pholidichthys. 











p. unguis 








SL 


69.1' 


76.7' 


77.32 


86.0' 


87.42 


1032 


2453 


Head length 


11.7(16.9) 


12.1 (15.8) 


12.1 (15.7) 


13.5 (15.7) 


13.6(15.6) 


14.6 (14.2) 


22.2 (9.1) 


Snout length 


2.2 (3.2) 


2.4(3.1) 


2.4 (3.1) 


2.8 (3.3) 


2.8 (3.2) 


3.0 (2.9) 


4.5 (1.8) 


Orbital diameter 


3.1 (4.5) 


2.5 (3.3) 


3.2 (4.1) 


3.0 (3.5) 


3.4 (3.9) 


3.2 (3.1) 


3.4 (1.4) 


Postorbital HL 


6.9 (10.0) 


7.6 (9.9) 


7.1 (9.2) 


8.2 (9.5) 


8.6 (9.8) 


9.0 (8.7) 


15.2 (6.2) 


Upper jaw length 


4.3 (6.2) 


4.1 (5.3) 


4.3 (5.6) 


4.6 (5.4) 


4.7 (5.4) 


5.3 (5.1) 


7.6 (3.1) 


Predorsal length 


11.9(17.2) 


12.5 (16.3) 


12.4(16.1) 


13.4(15.6) 


13.1 (15.0) 


14.2(13.7) 


25.1 (10.2) 


Body depth 


4.7 (6.8) 


5.0 (6.5) 


5.5 (7.1) 


5.9 (6.9) 


6.0 (6.9) 


6.3 (6.1) 


10.3 (4.2) 


Caudal-fin length 


5.1 (7.4) 


5.1 (6.6) 


5.2 (6.7) 


5.6 (6.5) 


5.5 (6.3) 


7.0 (6.8) 


14.5 (5.9) 


Mid-lateral dark 
















stripe depth 


0.6 (0.9) 


0.7 (0.9) 


0.5 (0.6) 


0.9 (1.0) 


1.0(1.1) 


0.9 (0.9) 


NA 










p. leucotaenia 








SL 


49.7^* 


52.15 


66.2^ 


77.65 


79.45 


80.2* 


83.3* 


Head length 


10.0(20.1) 


11.6(22.3) 


13.0(21.2) 


15.1 (19.5) 


15.4 (19.4) 


16.6 (20.7) 


16.4 (19.7) 


Snout length 


2.2 (4.4) 


2.5 (4.8) 


2.4 (3.9) 


3.2 (4.1) 


3.4 (4.3) 


3.4 (4.2) 


3.5 (4.2) 


Orbital diameter 


2.5 (5.0) 


3.0 (5.8) 


2.8 (4.6) 


3.0 (3.9) 


3.2 (4.0) 


3.2 (4.0) 


3.1 (3.7) 


Postorbital HL 


6.8 (13.7) 


7.0(13.5) 


8.2 (20.8) 


9.4(12.1) 


10.1 (12.7) 


10.2 (12.7) 


10.6 (12.7) 


Upper jaw length 


3.2 (6.4) 


3.8 (7.3) 


4.2 (6.8) 


5.2 (6.7) 


5.0 (6.3) 


5.3 (6.6) 


5.2 (6.2) 


Predorsal length 


11.8(23.8) 


10.7 (20.5) 


13.4(21.9) 


14.7 (18.9) 


15.3 (19.3) 


15.8 (19.7) 


15.5 (18.6) 


Body depth 


5.2 (10.4) 


4.6 (8.8) 


6.3 (9.5) 


6.7 (8.6) 


7.1 (8.9) 


8.8(11.0) 


9.3(11.2) 


Caudal-fin length 


4.2 (8.4) 


4.4 (8.4) 


4.3 (7.0) 


— 


— 


5.7 (7.1) 


6.9 (8.3) 


Mid-lateral dark 
















stripe depth 


3.0 (6.0) 


2.5 (4.8) 


3.3 (5.4) 


3.3 (4.3) 


2.8 (3.6) 


2.7 (3.4) 


2.7 (3.2) 










p. leucotaenia 








SL 


97.5^ 


102« 


115« 


224'° 


225'° 


337" 




Head length 


17.3 (17.8) 


18.9 (18.5) 


20.3 (17.6) 


30.5 (13.6) 


28.7 (12.8) 


41.8 (12.4) 




Snout length 


3.5 (3.6) 


3.6 (3.5) 


4.0 (3.5) 


6.3 (2.8) 


6.5 (2.9) 


7.8 (2.3) 




Orbital diameter 


3.5 (3.6) 


3.5 (3.4) 


3.6 (3.1) 


5.2 (2.3) 


4.6 (2.0) 


5.2 (1.5) 




Postorbital HL 


10.8(11.1) 


12.0(11.8) 


13.3 (11.6) 


20.5 (9.2) 


19.2 (8.5) 


31.0(9.2) 




Upper jaw length 


6.0 (6.2) 


6.0 (5.9) 


6.6 (5.7) 


10.9 (4.9) 


10.6 (4.7) 


14.1 (4.2) 




Predorsal length 


17.2 (17.6) 


18.2(17.8) 


18.7 (16.3) 


30.1 (13.4) 


27.8 (12.4) 


42.7 (12.7) 




Body depth 


8.5 (8.7) 


7.6 (7.4) 


8.9 (7.7) 


19.4 (8.7) 


18.1 (8.0) 


20.7 (6.1) 




Caudal-fin length 


6.5 (6.7) 


7.7 (7.5) 


8.9 (7.7) 


14.2 (6.3) 


14.4 (6.4) 


— 




Mid-lateral dark 
















stripe depth 


3.3 (3.4) 


2.8 (2.7) 


3.1 (2.7) 


NA 


NA 


NA 





'USNM 337860, 2NTM S. 13039-001, ^Holotype, NTM S. 11779-001; *AMS 1.34480-001, ^USNM 289924, 
6USNM 215258, ^AMS 1.34712-007, ^USNM 122340, "^USNM 150828, '"CAS 32048, "USNM 212163. 



describe the dark markings in their most 
pronounced and complete form. Tip of chin 
with dark smudge, another on snout above 
upper Up continuing ventrally below eye 
and posteriorly from mid-postorbital mar- 
gin as broad, dark stripe; stripe constricting 
at upper posterior margin of operculum and 
continuing as dark, slender midlateral stripe, 
which decreases in depth (depth at no point 
much greater than 1 mm, usually less) as it 
proceeds posteriorly to base of caudal fin, 
where it is briefly interrupted, beginning 



again, slightly intensified, on caudal fin, and 
extending, diffusely, for variable distance, 
up to the end of the fin. Slender, dark stripe 
dorsally on head, originating at postorbital 
margin, continuing posteriorly just ventral 
to dorsal-fin base, tapering and becoming 
fainter posteriorly until it vanishes, usually 
on or before reaching posterior third of 
body. No other prominent markings on 
head, body, or fins. 

Female adult (holotype, 245 mm SL). 
Head overall dusky with darker, diffuse 



VOLUME 109, NUMBER 2 357 

Table 2. — Frequency distributions for certain meristic characters of the species of Pholidichthys (localities 
arranged west to east), "x" denotes count recorded by Kailola (1973:11), but not encountered during our study. 
? denotes a specimen with a count equal to or slightly greater than indicated by the column heading. 

Dorsal-fin rays 
Species 66 67 68 69 70 71 72 73 74 75 76 77 78 79 . . . 86 87 88 89 90 91 92 93 94 95 96 97 98 

P. anguis 
J. Bonaparte Gulf 1 

N Melville Id 2 - - 1 

N. Goulbum Id 1' 3 112 11 

Cape Wessel 6 4 5 6 6-1-1 

Groote Eylandt 1 



P.leucotaenia 


XXXXXX331 18411 




Anal-fin rays 


Species 


49 50 51 52 53 54 55 56 57 58 59 60 61 62 . . . 70 71 72 73 74 75 76 77 78 79 80 81 



P. anguis 
J. Bonaparte Gulf ? 

N Melville Id 111 

N Goulbum Id 1' 3 - 1 3 2 1 1 ? 

Cape Wessel 2327452--1 

Groote Eylandt 1 



P. leucotaenia 


XXXXX21-443161 




Total vertebrae 


Species 


71 72 73 74 75 76 77 78 79 . . . 90 91 92 93 94 95 96 97 98 99 100 101 



P. anguis 
J. Bonaparte Gulf 1 

N. Melville Id 1-11 

N Goulbum Id 2 3 2 3 1 - ? 

Cape Wessel 18 9 3 7 3 1-11 

Groote Eylandt 1 



P. leucotaenia 


1-2432 


5 


6 1 




Precaudal vertebrae 




Caudal vertebrae 


Species 


22 23 24 25 26 27 


28 


48 49 50 51 52 53 54 55 56 . . . 64 65 66 67 68 69 70 71 72 73 74 75 



P. anguis 
J. Bonaparte Gulf 1 1 

N. Melville Id 1 1 11 

N. Goulbum Id 285 122412--? 

Cape Wessel 13 19 32 17886221 

Groote Eylandt 1 1 

P. leucotaenia 249111 217235311 

'Specimen has caudal vertebral fusions, which may have affected number of fin rays; caudal and total vertebral 
counts for this specimen not included in table. 

dusky blotch posterior to orbit; blotch in- at midlength, posteriorly) continuing as se- 

terrupted at preopercular margin, becoming ries of about 20 diffuse, dusky spots on 

much less distinct on opercle, continuing on body below midlevel, spots ending on pos- 

body as moderately large, indistinct series terior half of body. Series of diffuse ovoid 

of ovoid to elongate ovoid dusky areas, spots on dorsal body contour, beginning 

which become unrecognizable at about over posterior quarter of abdomen, continu- 

body midlength. Three horizontal pairs of ing to caudal-fin. 

dusky ovoid spots on abdomen (anteriorly. Dorsal and anal fins variably dusky, with 



358 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 






Fig. L Holotype of Pholidichthys unguis, NTM S.l 1799-001, female 245, mm SL, WSW of Groote Eylandt, 
Gulf of Carpentaria, Northern Territory, Australia. Bulge under lower jaw appears to be an abnormality. Full- 
length view is produced from two photographs pieced together. Arrows in lower two figures indicate same 
position in each photograph. (Photographs by T. B. Griswold) 



distal edge of each darker; both fins becom- 
ing much darker near posterior end, where 
they join darkly dusky caudal fin. Pectoral 
and pelvic fins pale dusky. A color photo- 
graph taken of the fresh specimen shows 
the spots on the body to be brownish; back- 
ground color tan. Large yellow eggs are 
visible through the transparent abdomen. 

Remarks. — The holotype has an appar- 
ently abnormal padlike swelling on the 
chin, possibly caused by a parasite. As seen 
through the body wall of the holotype (Fig. 
2), the maximum diameter of the eggs ap- 
pears to be approximately 2 mm. 

All specimens of P. unguis have come 



from benthic trawl samples from a mud 
substrate, which may include sand, shell, 
sponges, and rocky reef and coral patches. 
None have been found associated with coral 
reefs, as with P. leucotaenia. 

Comparisons. — Study of whole speci- 
mens and skeletal preparations of P. anguis 
indicates that the species exhibits the same 
familial specializations as P. leucotaenia 
(e.g., presence of a septal bone, fused 5th 
ceratobranchials, single nostril, no spinous 
fin rays, lack of scales, etc; see Springer & 
Freihofer 1976). Pholidichthys anguis dif- 
fers from P. leucotaenia in having 87-98 
dorsal-fin rays (vs. 66-79), 70-81 anal-fin 



VOLUME 109, NUMBER 2 



359 




Fig. 2. Holotype of Pholidichthys anguis, NTM S. 11799-001, from color photograph taken when specimen 
was freshly collected; note eggs visible through transparent abdominal skin. (Photograph by A. Baker) 



rays (vs. 49-62), 90-101 total vertebrae 
(vs. 71-79), a more slender body, shorter 
head, predorsal, and upper jaw lengths, 
lesser body depth (Fig. 6), and in the color 
pattern of juveniles, —50-103 mm SL 
(depth of slender dark stripe on midside at 
vertical from anal-fin origin up to about 1 
mm, usually much less, as opposed to depth 
2.5-3.3 mm in P. leucotaenia; Table 1; also 
compare Figs. 3 and 4). The adult color pat- 
tern, apparently, is also different from that 
of P. leucotaenia (Fig. 5), although appear- 
ing somewhat similar to that of the preadult 
(Figure 5, middle). Only one adult of P. 
anguis is known, and it is conceivable that 
the pattern is as variable as that of P. leu- 
cotaenia. Interestingly, adults of P. leuco- 
taenia are known only from aquarium 
raised juveniles, and none have been seen 
in the wild or collected. 

Etymology. — The specific name is from 
the Latin anguis, meaning "snake," refers 
to the elongate form of the species, and is 
here used as a noun in apposition. 



Distribution and Historical Biogeography 

Pholidichthys anguis is known only from 
the coastal waters of Northern Territory be- 
tween 128°56' and 136°15'E in depths of 
19-70 m (Fig. 7). Its distribution is allo- 
patric to that of P. leucotaenia, which is 
known from habitats close to or among cor- 
als, from the southern Philippines south to 
Flores, Indonesia, and east and south to 
New Caledonia. We question two other lo- 
cality records. Specimens indicated as orig- 
inating from Calatagan, Batangas [Prov- 
ince], Luzon, Philippines (CAS 32048, 
76415), were provided by aquarium im- 
porters, as seems to be the case with a spec- 
imen purportedly from Batangas (BMNH 
1982.8.3.4). A Batangas locality appears to 
be reasonable, but should be verified by 
better documented new collections. To ex- 
plain the current distributions of these two 
species, we propose the following scenario: 

Wilson & Allen (1987:64) concluded 
that "the pan-tropical Tethyan fauna and 
its successor, the modern Indo-West Pacific 



360 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 





Fig. 3. Pholidichthys anguis, juveniles. Northern Territory, Australia, a-c, NTM 13039-001, 87.8 mm SL, 
N of Melville Island, Arafura Sea, dusky markings relatively typical, c shows dorsal surface of head and 
anteriormost portion of body; d-e, NTM S. 13339-002, ca. 85, mm SL, Joseph Bonaparte Gulf, dusky markings 
unusually intense, e shows dorsal surface of head and anteriormost portion of body. (Photographs by T B. 
Griswold) 



fauna, have dominated the northern coast 
of Australia since the beginning of the Ter- 
tiary [ca. 65 m.y.a.]." We beheve their 
conclusion reasonable and hypothesize that 
the distribution of the common ancestor of 
the two species of Pholidichthys originated 
as part of the Tethyan fauna and occupied 
an area that excluded Australia-New Guin- 
ea. The ancestral distribution was possibly 
limited to the coastal areas of southeastern 
Asia (Malaya, Indonesia, Philippines) un- 
til, perhaps, as recently as mid-Miocene 
(ca. 16 m.y.a.), when southeast Asia and 
northern Australia-New Guinea were still 
separated by an expanse of deep, open sea. 
This expanse of sea acted as a barrier to 



dispersal between the two areas, particu- 
larly for shallow-dwelling marine organ- 
isms such as Pholidichthys, which lacks a 
planktonic stage (Trnski et al. 1989, Wirtz 
1991). After Australia-New Guinea collid- 
ed with the islands (Banda Arc) off south- 
east Asia (ca. 15 m.y.a.; Audley-Charles 
1981, 1987, Burrett et al. 1991), the spac- 
ing of existing and newly formed islands 
may have permitted shallow-dwelling or- 
ganisms to disperse between the colliding 
entities. If so, the common ancestor of the 
two extant Pholidichthys species was then 
able to disperse first to northern Australia- 
New Guinea and then into the New Brit- 
ain-New Ireland-Solomon volcanic islands 



VOLUME 109, NUMBER 2 



361 




Fig. 4. Pholidichthys leucotaenia juveniles (ca. 60 mm SL) photographed in aquarium (R. B. Hansen). 



chain. This chain originated along a mid- 
Pacific plate ridge during late Eocene-ear- 
ly Oligocene and was gradually displaced 
westward to converge with New Guinea 
during the past half million years (Kroenke 
1984, Yan & Kroenke 1993). After reach- 



ing the Solomons, Pholidichthys presum- 
ably dispersed south along the contiguous 
New Hebrides island chain (from which 
few fish collections have been made and 
Pholidichthys is as yet unreported) to 
close-by New Caledonia, and presumably 




v\^-n \ rrrv 1 




Fig. 5. Pholidichthys leucotaenia, aquarium specimens. Upper, adult, 205 mm SL, and middle, preadult, 108 
mm, purportedly originating from Philippines (from Springer & Freihofer 1976: fig. 23); bottom, ZMUC R75457, 
adult, ca. 175 mm SL, obtained from dealer in Singapore (G. Brovad, photographer). 



362 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




9 p. anguis y = 1.357x" 

irP. leucotaenia y = O.Veex"-**" 




- 








•k 


: / 


•k 


• 






^^ ^^^'^^ P- anguis 




y = 


6.667 


+ 0.0.075X 


jt^^^9f^ P- leucotaenia 


y = 


6.641 


+ 0.104X 

1 




P. anguis y = 3.069 + 

P. leucotaenia y = 0.2 Mx" 



SL (mm) 

Fig. 6. Scatter plots, best-fit curves, and regression 
formulae for certain mensurable characters in Pholi- 
dichthys species based on data listed in Table 1. 



the Loyalty Islands, with which the New 
Hebrides is colliding. 

The apparent absence of Pholidichthys 
from the Queensland coast and adjacent 
Great Barrier Reef, where one might expect 
it to occur, appears puzzling at first, but 



many marine forms appear to have been un- 
able to reach these areas from New Guinea, 
and vice- versa (a few didactic examples: 
the blenniid fishes Ecsenius namiyei, E. sel- 
lifer, E. taeniatus, E. trilineatus all occur in 
easternmost New Guinea, but are absent 
from the GBR; E. mandibularis, E. aus- 
tralianus, E. stictus all occur at the northern 
Queensland end of the GBR, but are absent 
from New Guinea — Springer (1988)). Coral 
reefs extend around the southeast tip of 
New Guinea and continue westward along 
its southern coast to about Port Moresby on 
the eastern side of the Gulf of Papua. The 
distribution of P. leucotaenia (Fig. 7) fol- 
lows this reefal distribution. A large num- 
ber of rivers drain into the western Gulf of 
Papua, which decreases salinity and in- 
creases turbidity in this portion of the Gulf, 
thus, creating a barrier to westward dis- 
persal of many marine forms that have 
reached the eastern portion of the Gulf. The 
most apparent (shortest) dispersal route to 
the Great Barrier Reef from New Guinea 
would be down the islands and reefs that 
extend north from the Cape York Peninsula. 
This route would be blocked to the north, 
however, by the riverine barrier in the west- 
ern portion of the Gulf of Papua, which we 
believe explains the absence of Pholidich- 
thys and many other apparently stenohaline 
forms from the Great Barrier Reef and 
Queensland. Pholidichthys is present in the 
western Gulf of Carpenteria, where small 
coral reefs are also present, but not in the 
eastern Gulf, where no reefs are present. 
Heavy river drainage and sediment deposi- 
tion in the eastern Gulf of Carpentaria prob- 
ably also prevents the eastward dispersal of 
reef obligates, such as Pholidichthys, 
around the coast of the Gulf of Carpentaria 
to eastern Queensland and the Great Barrier 
Reef (Wells 1957:pl. 9, charts the world 
distribution of coral reefs). 

The possibility of Pholidichthys' reach- 
ing Australia across the Arafura and Timor 
seas by dispersing along the southwestern 
coast of New Guinea and/or southeastern 
Indonesian island chain (Lesser Sunda to 



VOLUME 109, NUMBER 2 



363 




Fig. 7. Distribution of the species of Pholidichthys. Solid dots — P. leucotaenia; triangles — P. anguis; C = 
Gulf of Carpentaria; P = Gulf of Papua; shaded area represents emergent land joining northern Austraha and 
New Guinea during most recent glacial sea-level low, ca. 18,000 B.P. (other emergent land areas not shown). 



Am islands) would also present problems 
under present-day land-sea distributions. 
The southwestern coast of New Guinea 
bears numerous rivers draining into the 
Arafura sea and, consequently, the obligate 
coral-reef stepping stones are absent in the 
area. The present-day, relatively wide, cor- 
al-reefless expanse of the Timor and Ara- 
fura seas and the deep ocean paralleling and 
separating the southeastern islands from 
Australia are barrier enough to prevent the 
dispersal of forms such as Pholidichthys. 
Similarly, the barrier maintains the present- 
day allopatry of the two Pholidichthys spe- 
cies. 

During periods of Pleistocene glaciation 
(latest about 18,000 B.P.), the seaway pas- 
sage (Torres Strait) between central North- 
ern Territory and southern New Guinea was 
closed by a broad landbridge that extended 
between Northern Territory and much of 
southern New Guinea (Myers 1989:fig. 8, 
Springer & Williams 1990:fig. 7). The clo- 



sure of the passage probably would not 
have altered river flow into the western 
Gulf of Papua, but the emergent land would 
have eliminated ancestral Pholidichthys in 
the region from the Gulf of Carpentaria to 
somewhat west of Melville Island, ca. 
130°E. The present occurrence of Pholi- 
dichthys (P. anguis) in this relatively re- 
cently re-inundated area is the result of re- 
cent dispersal into Australian coastal habi- 
tats to the west. (Springer & Williams 1994: 
128, attribute east- west differences in the 
morphology of the blenniid Istiblennius 
meleagris, which is restricted to the north- 
ern coast of Australia, to the barrier created 
by formerly emergent land in the Gulf of 
Carpetaria-Melville Island region). We sug- 
gest that reef habitats formed along the 
western side of the Pleistocene landbridge 
that permitted dispersal of the ancestral spe- 
cies of Pholidichthys to northwestern Aus- 
tralia. As sea level rose and created modern 
conditions, the Australian and southeast 



364 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



Asian populations became isolated and di- 
verged. We are uncertain how, or if it is 
necessary, to accommodate for river runoff 
from the emergent west side of the land- 
bridge. Changes in the positions of river 
mouths on the west side would have been 
more radical than changes on the east side 
because of the greater area affected. Al- 
though we have chosen to base the final 
scene of our scenario on Pleistocene events 
of the past 18,000 years, a similar scenario 
could be developed for any of the earlier 
interglacials of the Quarternary. If only a 
small sample of the common ancestor 
reached Northern Territory and became iso- 
lated, rapid evolution (divergence) might be 
expected, as well as the current, apparently 
highly limited distribution of P. anguis. 

Acknowledgments 

For radiography, photography, routine 
laboratory assistance, and/or loan of speci- 
mens, we thank K. Murphy and J. Clayton 
(USNM), J. Nielsen (ZMUC), and D. Ca- 
tania (CAS). We extend our appreciation to 
A. Baker, R. Duckworth, and G. White, 
Northern Territory Fisheries Division, for 
bringing the first specimen to our attention. 
Suggestions for improvment of the manu- 
script were offered by R. Winterbottom 
(ROM) and R. D. Mooi (MPM). 

Literature Cited 

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the region of Wallace's Line. Pp. 24-35 in T. C. 
Whitmore, ed., Wallace's Line and plate tecton- 
ics, Clarendon Press, Oxford, 91 pp. 

. 1987. Dispersal of Gondwanaland: relevance 

to evolution of the angiosperms. Pp. 5-25 in T 
C. Whitmore, ed., Biogeographical evolution of 
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Bleeker, P. 1856. Bijdrage tot de kennis der ichthyol- 
ogische fauna van het eiland Boeroe. — Natu- 
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11:383-414. 

Burgess, W., & H. R. Axelrod. 1975. Pacific marine 
fishes vol. 6. T.F.H. Publications, Inc., Neptune, 
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Burrett, G., N. Duhig, R. Berry, & R. Varne. 1991. 
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ranes derived from Gondwana, and their bio- 
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Johnson, G. D. 1984. Percoidei: development and re- 
lationships. Pp. 464-498 in H. G. Moser, W. J. 
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832. 

Lockington, W N. 1882. List of the fishes collected 
by Mr. W J. Fisher upon the coasts of Lower 
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. 1988. The Indo-Pacific blenniid fish genus 

Ecsenius. — Smithsonian Contributions to Zool- 
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, & W C. Freihofer. 1976. Study of the mono- 

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niid fish genus Istiblennius reappraised: a revi- 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):366-372. 1996. 

A new species of Eleutherodactylus from Honduras related to 
Eleutherodactylus bransfordii (Anura: Leptodactylidae) 

Jay M. Savage, James R. McCranie, and Mario Espinal 

(IMS) University of Miami, Department of Biology, 

P.O. Box 249118, Coral Gables, Florida 33124-0421, U.S.A.; 

(JRM) 10770 SW 164th Street, Miami, Florida 33157-2933, U.S.A.; 

(ME) Residencial Cerro Grande, Bloque 48, 

Zona 2, Comayagiiela, Honduras 

Abstract. — A new species of Eleutherodactylus is described from northeast- 
ern Honduras. It is a member of the E. rhodopis group, differing from its closest 
congeners E. stejnegerianus of Pacific slope of Costa Rica and western Panama, 
by having definite toe webbing and a larger thenar tubercle, and E. bransfordii 
of Nicaragua to Panama, in lacking nuptial thumb pads in males and having a 
smaller thenar tubercle. 



A few years ago, the senior author (Sav- 
age 1981) referred a single specimen 
(LACM 45200) of small frog from north- 
eastern Honduras to Eleutherodactylus 
bransfordii (sensu Savage & Emerson 
1970) of Nicaragua, Costa Rica and Pana- 
ma. Subsequently, in 1986 and 1992-1994, 
examples of frogs of this general morph 
were collected by L. D. Wilson, J. R. 
McCranie and M. Espinal. Examination of 
these animals, plus a recently collected 
specimen in the Florida Museum of Natural 
History, showed several differences from 
typical E. bransfordii and led us to re-eval- 
uate the placement of the LACM specimen 
as well. Further study convinced us that the 
Honduran material was conspecific but rep- 
resented an undescribed species distinct 
from E. bransfordii. 

Methods 

All measurements are in millimeters and 
were made with dial calipers with the aid 
of a dissecting microscope. Abbreviations 
used are: SVL (standard length), HL (head 
length; tip of snout to angle of jaw), HW 
(head width; greatest width), SL (snout 
length; anterior border of eye to nostril), TL 
(tibia length; one end of tibia to other, in- 



cluding covering tissues), FL (foot length; 
distance from posteriormost portion of in- 
ner metatarsal tubercle to tip of fourth toe), 
TPL (tympanum length), EL (eye length), 
DW (third finger disc width). Snout outline 
and profile terminology follows Heyer et al. 
(1990), finger and toe disc, disc pad and 
subarticular tubercle shape terminology fol- 
lows Savage (1987), and color codes used 
for some of the color notes in life follow 
Smithe (1975). Museum acronyms follow 
Leviton et al. (1985). 

Eleutherodactylus lauraster, new species 
Figs. 1-2 

Holotype.—VSnm 344826, adult male, 
from above the Quebrada El Pifiol 
(15°07'N, 86°43'W), Parque Nacional La 
Muralla, Departamento de Olancho, Hon- 
duras, elevation 1200 m, collected 21 July 
1993 by Mario Espinal and James R. 
McCranie. Original field number LDW 
9997. 

Paratypes. — Honduras: Olancho: USNM 
344827-28, adult females, from the type lo- 
cality, 1180-1200 m; ROM 18095-96, 
adult males, 18097-98, adult females, from 
near the Rio Seco, in the Sierra de Agalta, 
NNW of Catacamas, 990-1000 m; UF 



VOLUME 109, NUMBER 2 



367 




Fig. 1. Adult female paratype of Eleutherodactylus lauraster (USNM 344830), SVL 19.7 mm. 



90217, adult female, from the Sierra de 
Agalta, ca. 9 km N Santa Maria del Real, 
1200 m; LACM 45200, adult female, from 
ca. 0.5 km SE San Jose del Rio Tinto, 330 
m; Gracias A Dios: USNM 344829-30, 
adult females, from the confluence of Que- 
brada Waskista and Rio Wampu, 85 m. 

Diagnosis. — A small species belonging 
to the Eleutherodactylus rhodopis group 
(Savage 1987), characterized by tiny males 
and slightly larger females that is distin- 
guished from its close allies, Eleutherodac- 
tylus bransfordii and E. stejnegerianus by 
having finger I definitely shorter than II and 
definite basal webbing between the toes. 
From the former, it further differs in having 
the thenar tubercle definitely smaller than 
the palmar tubercle and in lacking nuptial 
thumb pads in adult males (thenar and pal- 
mar tubercles about the same size and nup- 
tial pads present in adult male E. bransfor- 
dii). Although E. lauraster and E. stejne- 
gerianus lack male nuptial pads, the latter 
species has even smaller thenar tubercles 
than the Honduran form, being equal to or 



smaller than the basal subarticular tubercle 
on fingers II-III. 

Summary of characteristics. — Snout 
nearly rounded to rounded in dorsal aspect, 
rounded in profile; top of head flat; canthus 
rounded, distinct; loreal region concave; 
nostrils directed laterally, situated at a point 
about two-thirds distance between anterior 
border of eye and tip of snout; supratym- 
panic fold distinct, narrowly obscuring up- 
per edge of tympanum, tympanum other- 
wise prominent; narrow discs on fingers 
III-IV, with slightly pointed disc covers and 
swollen to cuspidate pads; no nuptial thumb 
pads; relative length of fingers I < II < IV 
< III; heels smooth to rugose, lacking tu- 
bercles; subarticular tubercles of hands and 
feet ovoid, projecting, usually obtuse in 
profile; six accessory palmar and numerous 
plantar tubercles; thenar tubercle definitely 
smaller than palmar tubercle, also some- 
what narrower and raised, larger than basal 
subarticular tubercles under fingers II-III; 
palmar tubercle ovoid; discs on all toes, 
those on toes III-IV with slightly pointed 



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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 





Fig. 2. Eleutherodactylus lauraster. Left: hand of adult female paratype (USNM 344828), scale 
right: foot of adult male holotype (USNM 344826), scale = 3 mm. 



1 mm; 



disc covers and swollen to cuspidate pads; 
inner metatarsal tubercle elliptical in out- 
line, projecting in profile; outer metatarsal 
tubercle rounded, projecting; relative length 
of toes I < II < V < III < IV; basal web- 
bing between toes; inner tarsal fold weak; 
skin on dorsal surface of body smooth to 
tuberculate (tubercles and/or rugosity usu- 
ally more obvious posteriorly), that of up- 
per eyelids rugose and tuberculate, that of 
belly and ventral surfaces of thighs coarsely 
areolate; a distinct small inguinal gland 
present; prevomerine tooth patches on ele- 
vated, somewhat triangular ridges between 
and behind choanae, each ridge separated 
by distance equal to or greater than size of 
either patch; vocal slits and sac absent. 

Coloration in life. — An adult female, 
USNM 344827, was recorded as follows: 
dorsal surfaces of head and body Dark Drab 
(color 119B) with Hair Brown (119A) 
markings: dorsal surfaces of limbs Army 



Brown (219B) with Sepia (219) crossbands; 
Sepia stripe extending from nostrils to eyes 
and continuing posteriorly from eyes to 
above point of insertion of forelimbs; pos- 
terior surfaces of thighs mottled pale brown 
and yellowish cream; ventral surfaces of 
head and body pale yellow with minute 
white flecks; ventral surfaces of thighs and 
groin pale yellow; iris copper with dark re- 
ticulations. Another adult female (USNM 
344828) was recorded as follows: dorsal 
surfaces of head and body Cinnamon-Drab 
(219C) with Sepia (219) markings; dorsal 
surfaces of limbs Vinaceous Pink (22 IC) 
with Sepia crossbands; iris gold with indis- 
tinct gold reticulations; rest same as that re- 
corded for USNM 344827. A third adult fe- 
male (USNM 344830) was recorded as fol- 
lows: dorsal surfaces of head and body Ma- 
hogany Red (132B) with a slightly darker 
middorsal hourglass-shaped figure and in- 
terocular bar; dorsal surfaces of limbs 



VOLUME 109, NUMBER 2 



369 



slightly paler than dorsum with dark Ma- 
hogany Red crossbands; posterior surfaces 
of thighs pale brown; all ventral surfaces 
and groin region flesh-colored; iris pale 
gold with black reticulations. 

Color in preservative. — Dorsal surface of 
body pale to dark brown with small, scat- 
tered darker brown spots or blotches pres- 
ent dorsally and dorsolaterally in all but 
darkest specimens; dorsal surfaces of limbs 
pale to medium-brown with darker brown 
crossbands; a distinct, dark brown facial 
and supratympanic stripe usually present; 
ventral surfaces of head and body pale 
cream-colored, with numerous, although 
usually widely separated, brown puncta- 
tions present on chin and throat, a few 
brown punctations may also be present on 
chest region and laterally on belly; ventral 
surfaces of thighs pale cream-colored, light- 
ly to heavily punctated with brown; poste- 
rior surfaces of thighs brown, although less 
densely punctated pale areas usually pres- 
ent. 

Measurements and proportions of holo- 
type.—SVL 17.9; HL 7.9; HL/SVL 0.441; 
HW 5.8; HW/SVL 0.324; SL 1.7; EL 2.2; 
SL/EL 0.773; TL 10.1; TL/SVL 0.564; FL 
9.1; FL/SVL 0.508; TPL 2.0; TPL/EL 
0.909; DW 0.2; DW/TPL 0.100. 

Measurements and proportions of para- 
types. — (Females following and separated 
from males by a comma; means in paren- 
theses) SVL 16.8-17.2 (17.0), 14.6-22.3 
(18.5); HL/SVL 0.413-0.417 (0.415), 
0.383-0.438 (0.411); HW/SVL 0.351- 
0.355 (0.353), 0.312-0.350 (0.331); SL/EL 
0.720-0.870 (0.795), 0.650-0.909 (0.780); 
TL/SVL 0.558-0.583 (0.571), 0.503-0.578 
(0.541); FL/SVL 0.517-0.530 (0.524), 
0.431-0.544 (0.488); TPL/EL 0.800-0.826 
(0.813), 0.500-0.682 (0.591); DW/TPL 
0.100-0.105 (0.103), 0.091-0.188 (0.140). 

Etymology. — The specific name pro- 
posed is a noun in apposition to honor our 
colleague Larry David Wilson. It derives 
from the Latin laurus = laurel, the source 
of the name Lawrence, and the diminutive 
suffix - aster, in allusion to his first name 



and his stature as herpetologist laureate for 
Honduras. 

Natural history notes. — Eleutherodacty- 
lus lauraster is known from between 85- 
1200 m in the Lowland Moist Forest and 
Premontane Wet Forest formations of Hold- 
ridge (1967). Specimens were collected on 
the forest floor, while active both during the 
day and at night. Two adult females with 
ovarian eggs (ROM 18098, USNM 344830) 
were collected 12 August and 28 July, re- 
spectively. 

Referred specimen. — Honduras: Gracias 
A. Dios: USNM 344831, juvenile, from the 
confluence of Quebrada Waskista and Rio 
Wampu, 85 m. 

Discussion 

Eleutherodactylus lauraster is assigned 
to the Eleutherodactylus rhodopis group 
{sensu Savage 1987). The characterization 
of the group as published by Savage (1987) 
now requires some modification. Firstly, a 
lapsus led to a statement implying that the 
members of this group have "an inner tarsal 
tubercle or two" (Savage 1987:49). The 
correct verbage is "an inner tarsal tubercle 
or two may be present," as this feature only 
occurs in Eleutherodactylus rhodopis. Sec- 
ondly, a weak inner tarsal fold is found in 
E. bransfordii, E. lauraster, E. podiciferus 
and E. stejnegerianus. Finally, the absence 
of toe webbing may no longer be used to 
define the group, since E. lauraster has bas- 
al webs on the toes and weak webs are 
sometimes present in some lower Central 
American forms. The remaining characters 
used by Savage (1987) to define the group 
remain valid, with the exception of the 
mandibularis muscle condition (see below). 

Lynch (1993) concluded that the depres- 
sor mandibularis muscle condition found in 
this group was misinterpreted by Savage 
(1987). Lynch (1993) generally rejected the 
systematic significance of differences in the 
characters of the depressor mandibularis 
utilized by Savage (1987) as modified from 
Starrett (1968). This is not the place to re- 



370 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



spond to Lynch's arguments, which will be 
addressed elsewhere. However, he did con- 
firm that examples of the five species of the 
E. rhodopis group that he examined share 
the DFSQdAT (as pointed out by Starrett, 
1968, contra Savage 1987). This feature is 
unique among Eleutherodactylus belonging 
to the Middle American clade (sensu Lynch 
1986) or "subgenus Craugaster." 

All other members of the Craugaster clade 
diagnosed by the synapomorphy of having 
only the externus superficialis adductor 
mandibularis (e) present, have dfsq, 
DFsqat, or DFSQAT conditions of the ad- 
ductor. The jaw muscle formula for the E. 
rhodopis group is corrected to DFSQdAT 
+ e and is diagnostic. Lynch (1993) re- 
ported this formula for Eleutherodactylus 
bransfordii, E. hobartsmithi, E. mexicanus, 
E. pygmaeus, and E. rhodopis. We add E. 
lauraster, E. podiciferus, and E. stejneger- 
ianus to that list. We predict that the re- 
maining species of the E. rhodopis group 
(£". jota, E. saltator, and E. sartori) also 
have the same jaw muscle configuration. 

The status of populations placed in Eleu- 
therodactylus bransfordii Cope (1886) by 
Savage & Emerson (1970) are in a state of 
flux. Miyamoto (1983) pointed out that in 
Costa Rica, Pacific slope samples differ sig- 
nificantly in allozyme features from Atlan- 
tic versant frogs and revived the name Eleu- 
therodactylus stejnegerianus Cope (1893) 
for them. Morphological features have now 
been determined to distinguish between 
these forms as well (see diagnosis of Eleu- 
therodactylus lauraster and the key below). 

Eleutherodactylus bransfordii and E. 
stejnegerianus are essentially allopatric but 
overlap geographically at a few sites along 
low passes near the continental divide in 
northern Costa Rica. In addition, Miyamo- 
to's (1983) allozyme evidence indicated 
that a third cryptic species occurred with E. 
stejnegerianus at some Costa Rican locali- 
ties. The status of this form and its relations 
to the other recognized species continue un- 
der investigation. 

Eleutherodactylus lauraster is somewhat 



intermediate in characters between E. 
bransfordii and E. stejnegerianus but dif- 
fers from both in having a short thumb. It 
most closely resembles the latter species in 
having a reduced thenar tubercle and in 
lacking nuptial pads in adult males. How- 
ever, neither of these features is conclusive 
evidence of close relationship. Because on- 
going work on the karyology of this group 
by S. H. Chen gives promise of resolving 
the phylogeny of the cluster of bransfordii- 
like species, any conclusions based on the 
few morphological features separating the 
recognized forms remain premature. 

The recognition of Eleutherodacylus 
lauraster brings the number of species re- 
ferred to the E. rhodopis group to eleven. 
The following key may be used to identify 
the included taxa. 

Key to the Frogs of the Eleutherodactylus 
Rhodopis Group 

la. No inner tarsal tubercles 2 

lb. One or two distinct inner tarsal tu- 
bercles (Atlantic slope: San Luis 
Potosi, Mexico, to western Hon- 
duras; Pacific slope: Oaxaca, 

Mexico, to El Salvador) 

E. rhodopis 

2a. A thenar and 1 palmar tubercle; 
nuptial pads and/or vocal slits 
present or not in adult males ... 3 

2b. A thenar and 2 palmar tubercles 
that may be partially fused; no 
nuptial pads or vocal slits in adult 
males 9 

3a. No inner tarsal fold; no vomerine 
teeth; no nuptial pads or vocal 
slits in adult males 4 

3b. A definite inner tarsal fold; vo- 
merine teeth present 5 

4a. A row of tubercles along outer 
edge of tarsus (Central Mexico 
from Colima to Guerrero, and the 
state of Mexico) . . . E. hobartsmithi 

4b. No row of tubercles on outer edge 
of tarsus (Atlantic slope: tropical 



VOLUME 109, NUMBER 2 



371 



Mexico; Pacific slope: Jalisco, 

Mexico, to Guatemala) 

E. pygmaeus 

5a. No enlarged calcar on heel; no ul- 
nar or outer tarsal folds 6 

5b. An enlarged triangular calcar on 
heel; ulnar and outer tarsal folds 
present (western Panama) . . . E. jota 

6a. Heel smooth to granulate; tuber- 
cles on underside of hand pro- 
jecting 7 

6b. One to 3 enlarged heel tubercles; 
tubercles on undersides of hands 
low and rounded; no nuptial pads 
but vocal slits present in adult 
males (uplands of Costa Rica and 
western Panama) E. podiciferus 

7a. Thenar and palmar tubercles 
about same size, larger than sub- 
articular tubercles on fingers II- 
III; nuptial pads present but no 
vocal slits in adult males (Atlantic 
slope from Nicaragua to central 
Panama) E. bransfordii 

7b. Thenar tubercle much smaller 
than palmar tubercle, no nuptial 
pads or vocal slits in adult males 
8 

8a. Definite basal toe webbing pres- 
ent; finger I definitely shorter than 
finger 11; thenar tubercle larger 
than basal subarticular tubercle 
under finger III (Atlantic slope of 
eastern Honduras) E. lauraster 

8b. No toe webbing or slight webs 
present basally only between toes 
II-IV; fingers I and II equal; the- 
nar tubercle equal to basal subar- 
ticular tubercle under finger III 
(Pacific slope from Costa Rica to 
western Panama) . . E. stejnegerianus 

9a. Inner metatarsal tubercle much 
larger than outer 10 

9b. Metatarsal tubercles subequal in 
size (Pacific slope of Chiapas, 

Mexico) E. sartori 

10a. Outer edge of tarsus with row of 
small tubercles (uplands of south- 



ern Mexico, west of Isthmus of 

Tehuantepec) E. mexicanus 

10b. Outer edge of tarsus smooth (up- 
lands of Guerrero and western 
Oaxaca, Mexico) E. saltator 



Acknowledgments 

Field assistance was provided by G. A. 
Flores, J. Rindfleish, K. L. Williams, and L. 
D. Wilson. Expert service as guides and 
collectors was provided by D. Almendarez 
of El Dictamo, Olancho, and E. Flores of 
La Llorona, Olancho. Collecting and ex- 
portation permits were provided by W. 
Aguilar, formerly of the Departamento de 
Recursos Naturales Renovables, Tegucigal- 
pa, and by E. Munoz and G. Rodriguez of 
COHDEFOR, Tegucigalpa. We are also 
grateful to the following institutions and cu- 
rators for the loan of material: R. L. Bezy 
(LACM); R. W Murphy (ROM); D. L. 
Auth (UF). Jay M. Savage extends partic- 
ular thanks to the John Simon Guggenheim 
Foundation and Organization for Tropical 
Studies (OTS) for support for studies of 
Central American amphibians. 

Literature Cited 

Cope, E. D. 1886. Thirteenth contribution to the her- 
petology of tropical America. — Proceedings of 
the American Philosophical Society 23:271- 
287. 

. 1893. Second addition to the knowledge of 

the Batrachia and Reptilia of Costa Rica. — Pro- 
ceedings of the American Philosophical Society 
31:333-347. 

Heyer, W. R., A. S. Rand, C. A. G. da Cruz, O. L. 
Peixoto, & C. E. Nelson. 1990. Frogs of Bor- 
aceia. — Arquivos de Zoologia, Museu de Zool- 
ogia, Universidade de Sao Paulo 31:231-410. 

Holdridge, L. R. 1967. Life zone ecology. Revised ed. 
Tropical Science Center, San Jose, Costa Rica, 
206 p. 

Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. 
Dawson. 1985. Standards in herpetology and 
ichthyology: Part I. Standard symbolic codes 
for institutional resource collections in herpe- 
tology and ichthyology. — Copeia 1985:802- 
832. 

Lynch, J. D. 1986. The definition of the Middle 



372 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



American clade of Eleutherodactylus based on 
jaw musculature (Amphibia: Leptodactyli- 
dae).— Herpetologica 42:248-258. 
— . 1993. The value of the m. depressor man- 



dibulae in phylogenetic hypotheses for Eleu- 
therodactylus and its allies (Amphibia: Lepto- 
dactylidae). — Herpetologica 49:32-41. 

Miyamoto, M. M. 1983. Biochemical variation in the 
frog Eleutherodactylus bransfordii: Geographic 
patterns and cryptic species. — Systematic Zo- 
ology 32:43-51. 

Savage, J. M. 1981. Eleutherodactylus bransfordii 
(Cope): An addition to the frog fauna of Hon- 
duras. — Herpetological Review 12:14. 

. 1987. Systematics and distribution of the 



Mexican and Central American rainfrogs of the 
Eleutherodactylus gollmeri group (Amphibia: 
Leptodactylidae). — Fieldiana: Zoology, New 
Series 33:i-iv, 1-57. 
— , & S. B. Emerson. 1970. Central American 



frogs allied to Eleutherodactylus bransfordii 
(Cope): a problem of polymorphism. — Copeia 
1970:623-644. 

Smithe, F. B. 1975. Naturalist's color guide. Part I. 
Color guide. American Museum Natural His- 
tory, New York, 182 color swatches. 

Starrett, P. A. 1968. The phylogenetic significance of 
the jaw musculature in anuran amphibians. Un- 
published Ph.D. Dissertation, University of 
Michigan, Ann Arbor, 179 pp. 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):373-390. 1996. 

Hybrid wood warblers, Dendroica striata x Dendroica castanea 

(Aves: Fringillidae: Tribe Parulini) and the diagnostic 

predictability of avian hybrid phenotypes 

Gary R. Graves 

Department of Vertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — The plumage pattern and color and external morphology of hy- 
brid wood warblers {Dendroica striata X Dendroica castanea) are described. 
This hybrid combination constitutes the only known case of hybridization be- 
tween two broadly sympatric species of the genus Dendroica (Fringillidae; 
Tribe Parulini), that is represented by both male and female specimens in de- 
finitive alternate plumage. The plumage of the hybrids exhibited a mosaic of 
character states that varied in the degree of intermediacy between those of the 
parental species. External measurements of the hybrids fell within the cumu- 
lative ranges of characters of the postulated parental species. I hypothesize that 
the diagnostic predictability of hybrid phenotypes decreases as an inverse func- 
tion of genetic relatedness of the hybridizing species. 



Plumage color patterns are remarkably 
diverse among the 24 species of Dendroica 
(Aves: Fringillidae; Tribe Parulini) of North 
America and the Caribbean (taxonomy of 
Sibley & Monroe 1990), in contrast to the 
similarity of their body plans and trophic 
appendages. Several pairs of species 
thought to have recently diverged (Mengel 
1964, Mayr & Short 1970, Bermingham et 
al. 1992) exhibit striking differences in 
male definitive plumage, suggesting that 
plumage pattern may evolve rapidly in re- 
sponse to sexual selection. Our understand- 
ing of the inheritance patterns of plumage 
color in Dendroica has been limited to ev- 
idence gleaned from hybridization at con- 
tact zones between parapatric taxa (e.g., 
Dendroica occidentalis X Dendroica town- 
sendi; see Rohwer 1994) and rare instances 
of intergeneric hybridization (e.g., Parkes 
1978, Graves 1993a). 

Brodkorb's (1934) description of the hy- 
brid, Dendroica striata (Blackpoll Warbler) 
X Dendroica castanea (Bay-breasted War- 
bler), was the first report of hybridization 



between broadly sympatric species of Den- 
droica. This specimen (University of Mich- 
igan Museum of Zoology [UMMZ] No. 
53692) represents one of the few known in- 
trageneric Dendroica hybrids represented 
by a male in definitive alternate plumage. 
Because male D. striata and D. castanea 
differ dramatically in plumage pattern and 
color, this specimen assumes unusual sig- 
nificance for the investigation of phenotypic 
inheritance in avian hybrids. 

Recently, the existence of a female hy- 
brid (UMMZ 216628) and an additional 
male hybrid (San Bernardino County Mu- 
seum, uncataloged) identified as D. striata 
X D. castanea, was brought to my atten- 
tion, respectively, by Janet Hinshaw and 
Kenneth Parkes. Here I perform a hybrid 
diagnosis on all three specimens based on 
plumage color and pattern and external 
morphology, following Graves (1990). 

Materials and Methods 

The male specimen reported by Brod- 
korb (1934) was collected on 19 May 1920, 



374 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 










Fig. L Lateral view (from top) of males: Dendroica castanea; hybrids, D. striata X D. castanea (UMMZ 
53692 and SBCM); and D. striata (bottom). 



by Norman A. Wood at Warren Dunes, Ber- 
rien County, Michigan (Figs. 1-4). The fe- 
male specimen was collected on 25 May 
1970 at Long Point, Ontario, by Joseph G. 
Strauch (Figs. 5-7). The second male spec- 
imen was collected by Lawrence Sansone, 
III, along the Rio Grande south of Browns- 
ville, Cameron County, Texas, on 9 June 
1969. For the purposes of the hybrid diag- 
nosis, I considered all wood warblers that 
regularly breed north of the Mexican border 
as potential parental species {n = 53). 

The first two specimens have broadly ta- 
pered rectrices indicating that they were 
adults in their second year or older (ASY 
in banding terminology) (Pyle et al. 1987). 
Plumage, especially the rectrices, of the 
Texas specimen is moderately worn. Thus, 
I am uncertain of the bird's age (SY or 
ASY). Because the specimens appeared to 
be in definitive plumage, I compared them 



to large series of definitive-plumaged spec- 
imens in the collections of the National Mu- 
seum of Natural History, Smithsonian In- 
stitution. 

For comparative purposes I measured the 
size of three species: D. striata, D. casta- 
nea, and D. pinus. Data for D. pinus were 
omitted from Table 1, after analyses 
showed that it was not involved in the par- 
entage of the hybrids. Measurements of 
wing chord, tail length (from point of in- 
sertion of central rectrices to tip of longest 
rectrix), tarsus length, and bill length (from 
anterior edge of nostril), were made with 
digital calipers to the nearest 0.1 mm. 

I evaluated the color of selected areas of 
plumage with a Color Mate Colorimeter 
(Milton Roy), employing a 9.4 mm aper- 
ture. The data in Table 2 were compiled 
from the averages of three independent 
measurements (specimen removed from ap- 



VOLUME 109, NUMBER 2 



375 



rmted stutes "^O^^ 7^ N-,u )i • \ 



Locality A 




Cnll'i"r , 



(S"/ 



Fig. 2. Dorsal view (from top) of males: Dendroica castanea; hybrids, D. striata X D. castanea (UMMZ 
53692 and SBCM), and D. striata (bottom). 



erture between trials) per specimen for each 
plumage area. 

Colorimetric characters were described in 
terms of opponent-color coordinates (L, a, 
b) (Hunter & Harold 1987). This system is 
based on the hypothesis that signals from 
the cone receptors in the human eye are 
coded by the brain as light-dark (L), red- 
green (fl), and yellow-blue {b). The ration- 
ale is that a color cannot be red and green 
or yellow and blue at the same time. There- 
fore, "redness" and "greenness" can be 
expressed as a single value a, which is pos- 
itive if the color is red and negative if the 
color is green. Likewise, "yellowness" or 
"blueness" is expressed by b for yellows 
and —b for blues. The third coordinate L, 
ranging from to 100, describes the "light- 



ness" of color; low values are dark, high 
values are light. In other words, the more 
light is reflected from the plumage the high- 
er the L value will be. 

Difference among character means of 
presumed parental species were evaluated 
with two sample t-tests (Wilkinson 1989). 
Significance of probability values was ad- 
justed for the number of simultaneous tests 
of morphology (Table 1; P = 0.05/8 = 
0.006) and plumage color (Table 2; P = 
0.05/12 = 0.004). 

Hybrid diagnoses followed a two-step 
procedure. First, the presumed parental spe- 
cies of each hybrid were hypothesized 
through the visual comparison of plumage 
pattern and color. These hypotheses were 
then examined with quantitative analyses of 



376 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




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Fig. 3. Ventral view (from top) of males: Dendroica castanea; hybrids, D. striata X D. castanea (UMMZ 
53692 and SBCM); and D. striata (bottom). 



colorimetric and morphometric data. Con- 
cordance of results was interpreted as 
strong support for the presumed parentage 
of the hybrid (Graves 1990, 1993a; Graves 
& Zusi 1990). 

I used principal components analysis 
(PC A) on log 10 transformed variables to re- 
duce the dimensionality of data and to fa- 
cilitate the analysis of morphology in two 
dimensions. Unrotated principal components 
were extracted from covariance matrices 
(Wilkinson 1989). Because the external 
measurements of D. striata and D. castanea 
overlap (Table 1), both species and sexes 
were pooled for PCA. Separate PCA for 
each sex were performed on logjo trans- 
formed colorimetric variables of back plum- 
age (Tables 3, 4). Bivariate plots of factor 
scores from PCA analyses that included both 
back and crown color were less informative 



because significant differences in crown col- 
or of the parental species of both sexes po- 
larized the distribution of factor scores. 

Results 

Plumage Characters 

Males. — I concur with Brodkorb's (1934) 
identification but his brief, one-paragraph 
description of the Michigan specimen did 
not address alternate hypotheses or the ex- 
ternal morphology of the parental species 
(p. 243): 

"It is similar to D. striata, but differs 
from the latter species in the following 
particulars: malar region and chin black, 
with only the anterior half of the inter- 
ramal region and a spot one or two mil- 
limeters in length at the end of the rami 



VOLUME 109, NUMBER 2 



377 





Fig. 4. 
(bottom). 



Lateral view of male hybrids, Dendroica striata X D. castanea: UMMZ 53692 (top) and SBCM 



white; black streaks on the sides of throat 
coalescent, leaving a white line in the 
center of the throat only one to two mm. 
broad; chin, malar region, sides of neck, 
sides and flanks more or less marked with 



bay; suborbital, auricular, and postauri- 
cular regions, as well as breast, pale 
cream buff; the abdomen, sides, flanks, 
and under tail-coverts also somewhat suf- 
fused with this color." 



378 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




^^^ 




Fig. 5. Lateral view of female Dendroica castanea (top), hybrid, D. striata X D. castanea (UMMZ 216628), 
and D. striata (bottom). 



Hybrid wood warblers can be notoriously 
difficult to identify (Short & Robbins 1967, 
Graves 1988). In this case, however, iden- 
tification was expedited by the distinctive 
characters of the Michigan specimen: (1) 
black crown (Fig. 1); and (2) mixture of 
black and chestnut in the malar region (Fig. 
2). The black crown of the hybrid, which 
extends ventrally to the lower margin of the 
eye, is shared with Dendroica striata and 
the morphologically distinctive Setophaga 
ruticilla. The latter species can be conclu- 
sively excluded as a parental species for 
several reasons, most notably because the 
rectrices and remiges of the hybrid lack or- 
ange or yellow spots, even as traces. There- 
fore, by default, D. striata was one of the 
parental species. Determination of the other 



parental species was also relatively simple. 
Chestnut and black feathers on upper throat 
and malar regions of the hybrid could have 
been contributed only by Dendroica cas- 
tanea. 

Although the plumage of the Texas spec- 
imen differs in detail from the Michigan 
male, its diagnostic characters are the same 
(Appendix 1). In sum, plumage characters 
of these male hybrids can only be account- 
ed for by two species, D. striata and D. 
castanea. Other pairs of warbler species 
lack the range of pattern elements and 
plumage colors exhibited by the hybrids. 

Female. — Plumage characters of the hy- 
brid that were critical to its identification 
can be categorized as color or pattern ele- 
ments, although these are not mutually ex- 



VOLUME 109, NUMBER 2 



379 



''^^cr 



fS// 








Fig. 6. Dorsal view of female Dendroica castanea (top), hybrid, D. striata X D. castanea (UMMZ 216628), 
and D. striata (bottom). 



elusive. Prominent pattern elements in- 
clude: (1) streaked crown, mantle, and 
scapulars; (2) poorly defined superciliary; 
(3) a well-defined neck patch; (4) muted 
streaks on the flanks and sides of breast; 
and (5) contrasting tips of the middle and 
greater wing coverts that form wing bars. 
Important color elements include: (1) olive 
crown and olive-gray mantle with blackish- 
brown shaft streaks; (2) buffy neck patch; 

(3) buff throat, upper breast, and flanks; and 

(4) pale chestnut markings on flanks. 

Of the (^2^) = 1378 possible pairwise 
combinations of wood warbler species in 
the geographic species pool, the number 
that could have produced each of the afore- 
mentioned hybrid characters was substan- 
tial. For example, 1027 pairs of species 



could have produced a hybrid with contrast- 
ing wing bars because one or both of the 
species possessed them. Of the many pos- 
sible color and pattern characters present in 
the hybrid, only one — the buffy post-auric- 
ular neck patch — is restricted to a single 
species. Although females of several other 
species of Dendroica have a tendency to 
show a faint post-auricular patch in basic 
plumage, only D. castanea has a well-de- 
veloped buff patch in definitive alternate 
plumage. Other characters of the hybrid, 
which are shared with D. castanea and two 
other species, are a buff throat and breast 
(shared with Helmitheros vermivorus) and 
chestnut markings on the flanks (also 
shared with Dendroica pensylvanica). Hy- 
bridization between the latter two species. 



380 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 




Fig. 7. Ventral view of female Dendroica castanea (top), hybrid, D. striata X D. castanea (UMMZ 216628), 
and D. striata (bottom). 



Table 1. — Ranges and means (± one standard deviation) of measurements of definitive-plumaged males and 
females collected in May and June of Dendroica castanea, D. striata, and their putative hybrids (University of 
Michigan Museum of Zoology, No. 53692, 3; No. 216628, ?; San Bernardino County Museum (SBCM), S). 
An asterisk indicates that character means of D. striata and D. castanea differed at F = 0.006 (= 0.05/8). 







striata 


castanea 




Hybrids 






UMMZ 






66 
(n = 12) 


95 
(n = 12) 


66 
(n = 12) 


99 
(n = 12) 




Character 


6 


9 


6 


Wing 


71.0 - 76.8 


68.6 - 70.7 


71.0 - 75.9 


68.0 - 72.1 


71.3 


68.8 


70.1 


chord 


73.9 ± 1.9 


70.7 ± 1.4 


73.2 ± 1.5 


70.6 ± 1.1 








Tail 


47.6 - 55.2 


47.4 - 50.9 


51.0 - 55.6 


48.7 - 52.3 


50.3 


49.5 


50.1 




51.3 ± 2.0 


*49.8 ± 1.0 


52.5 ± 1.4 


51.0 ± 1.0 








Tarsus 


18.1 - 19.9 


17.5 - 18.7 


16.5 - 19.1 


16.8 - 18.2 


17.5 


18.5 


18.2 




*18.9 ± 0.6 


18.1 ± 0.4 


17.8 ± 0.8 


17.6 ± 0.5 








Bill 


7.7 - 8.8 


7.6 - 8.5 


7.3 - 8.7 


7.4 - 8.4 


7.7 


7.7 


8.3 




8.1 ± 0.3 


8.0 ± 0.3 


8.1 ± 0.4 


7.9 ± 0.3 









VOLUME 109, NUMBER 2 



381 



Table 2. — Spectrophotometric measurements of back and crown color of male and female Dendroica striata, 
D. castanea, and their hybrids. Asterisks indicate character means of D. striata and D. castanea are significantly 
different at P = 0.004 (= 0.05/12). 







Males 














Hybrid 




striata 
(n = 12) 




castanea 
(n = 12) 






(UMMZ) 


(SBCM) 


Back 




















Lightness (L) 


26.5 - 32.9 
29.2 ± 2.1 




28.8 
31.3 


+ 


37.0 

2.5 






29.4 


33.6 


Red {a) 


0.3 - 1.2 
0.8 ± 0.3 




0.8 
1.4 


-H 


3.0 
0.6 






1.5 


1.4 


Yellow ib) 


4.7 - 11.1 
7.2 ± 1.8 




7.1 
8.8 


+ 


13.1 
1.6 






7.9 


8.5 


Crown 




















Lightness (L) 


13.0 - 16.9 

*14.4 ± 1.4 




20.0 

22.5 


+ 


25.0 
1.6 






16.9 


14.4 


Red (a) 


0.2 - 0.9 
*0.6 ± 0.2 




8.7 
12.2 


-H 


14.5 
1.8 






1.5 


2.4 


Yellow (J?) 


0.2 - 2.1 
*1.1 ± 0.5 




9.5 
13.6 

Females 


+ 


16.6 
2.0 






2.4 


4.5 




striata 
(n = 12) 






castanea 
(n = 12) 




Hybrid 
(UMMZ) 


Back 




















Lightness (L) 


28.5 - 


33.7 






29.9 


- 


34.3 




33.8 




31.6 ± 


1.7 






32.6 


+ 


1.3 






Red (a) 


0.8 - 


2.6 






0.7 


- 


1.5 




1.4 




1.6 ± 


0.6 






1.2 


-H 


0.2 






Yellow {b) 


9.1 - 


14.8 






7.5 


- 


11.6 




9.0 




*12.4 ± 


1.9 






9.1 


-¥■ 


1.1 






Crown 




















Lightness (L) 


24.5 - 


34.1 






24.3 


- 


32.4 




29.8 




28.9 ± 


3.1 






28.0 


-+- 


2.8 






Red (a) 


0.5 - 


2.5 






1.0 


- 


12.9 




1.4 




*1.4 ± 


0.6 






8.0 


■+- 


3.8 






Yellow {b) 


10.0 - 


18.3 






11.8 


- 


18.1 




12.6 




13.5 ± 


2.3 






14.7 


+ 


2.0 







Table 3. — Factor loadings for the first two principal 
components from an analysis of back color in male 
Dendroica striata, D. castanea, and the male hybrids 
(UMMZ 53692 and SBCM). 



Table 4. — Factor loadings for the first two principal 
components from an analysis of back color in female 
Dendroica striata, D. castanea, and the female hybrid 
(UMMZ 216628). 





Principal component axes 


Variable 


Principal ( 


;omponent axes 


Variable 


I 


II 


I 


II 


Lightness (L) 


0.007 


0.021 


Lightness (L) 


0.004 


0.002 


Red (a) 


0.193 


-0.025 


Red (a) 


0.137 


0.026 


Yellow (b) 


0.076 


0.062 


Yellow (b) 


0.049 


-0.073 


Variance explained 


88.4% 


10.0% 


Percent variance explained 


76.4% 


21.9% 



382 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



CM 

O 
Q_ 



0,1 



0.0 - 



-0,1 - 



-0,2 - 



-0.3 







1 




U 1 




- 






o 


o \ 


- 




o 






• • \ 

o • 


• 






\o 


o 


o • / 








1 


o 


o y 

1 





1.0 



-0.5 



0.0 



0.5 



PC 1 



Fig. 8. Bivariate plots of factor scores from a principal components analysis of back color (see Table 3) for 
male Dendroica striata (empty circles), D. castanea (filled circles), and D. striata X D. castanea hybrids 
(Michigan hybrid = diamond; Texas hybrid = square). The curved line in this and other plots represents the 
95% confidence ellipse. 



however, could not have produced the other 
characters present in the hybrid. Thus, D. 
castanea is clearly indicated as one of the 
parental species. 

The olive crown and muted flank streaks 
of the hybrid could not have been contrib- 
uted by D. castanea and must be characters 
of the other parent. Among female wood 
warblers in definitive alternate plumage, the 
only species possessing both yellow or ol- 
ive crowns and lateral ventral streakings are 
D. striata and D. pinus. The eye ring, chin, 
throat, breast, and upper belly of D. pinus 
vary from dull to bright yellow, and its dor- 
sal plumage is heavily saturated with yel- 
low. The hybrid has little yellow in the un- 
derparts especially the breast, exhibiting 
about the same amount of yellow as ex- 
pressed in series of D. striata, but consid- 
erably less than that of the dullest, defini- 
tive-plumaged D. pinus. This suggests that 
D. pinus is highly unlikely to have been a 



parental species. Thus, by the process of 
elimination, the second parental species was 
D. striata. 

In sum, the plumage pattern and color of 
the female hybrid can be accounted for by 
the two most probable parental species, 
Dendroica striata and D. castanea (Appen- 
dix 2). Other pairs of species lack the di- 
versity of plumage color and pattern ele- 
ments exhibited by the hybrid. 

Colorimetric Variation 

Univariate comparisons. — Male D. stria- 
ta are significantly darker than D. castanea 
on the back and crown, while the back and 
crown of D. castanea are both redder and 
yellower than those of D. striata (Table 2). 
As Brodkorb (1934) implied in his descrip- 
tion, the crown of the Michigan hybrid is 
similar in general appearance to that of D. 
striata (see Appendix 1). Several of the 



VOLUME 109, NUMBER 2 

0.2 
0,1 



383 



CM 

O 
Q_ 



0.0 - 



-0.1 - 



-0.2 




-0.3 -0,2 -0.1 



PC 1 



Fig. 9. Bivariate plots of factor scores from a principal components analysis of back color (see Table 4) for 
female Dendroica striata (empty circles), D. castanea (filled circles), and a D. striata X D. castanea hybrid 
(diamond). 



crown feathers of both male hybrids have 
buffy-olive tips or margins. These cause the 
colorimetric values for red {a) and yellow 
{b) to fall outside the range of values ex- 
pressed by ASY males of D. striata. Back 
color of the Michigan hybrid is intermedi- 
ate between that of D. striata and D. cas- 
tanea, while values for the Texas hybrid 
were closer to the characters means of D. 
castanea. 

Colorimetric variables of female D. stria- 
ta and D. castanea overlap (Table 2). Back 
color of the female hybrid more closely re- 
sembles that of D. castanea. The colori- 
metric values for lightness and redness of 
the hybrid fall narrowly outside the char- 
acter ranges for D. striata. Crown color 
variables of D. striata and D. castanea also 
overlap, but those of D. castanea are sig- 
nificantly redder and slightly yellower. 
Back color of the hybrid approximates the 
character means of D. striata. 

Multivariate comparisons. — ^Factor scores 



from a principal components analysis of 
back color of male D. striata and D. cas- 
tanea overlapped slightly along the first 
axis, which largely reflected the difference 
in redness between the species (Fig. 8). The 
distribution of D. castanea along the second 
PCA axis, which explained only a tenth of 
the variance, was entirely overlapped by D. 
striata. The male hybrids most closely re- 
sembled D. castanea in bivariate space and 
occurred well within the 95% confidence 
ellipse for hybrids and parental species. 

Factor scores for female back color of D. 
striata and D. castanea overlapped moder- 
ately along the first axis (PCA 1) and ex- 
tensively along the second (Fig. 9, Table 4). 
Factor scores in the upper left hand quad- 
rant represent specimens that are redder 
than average. The female hybrid most 
closely resembled D. castanea in color and 
fell narrowly outside the zone of overlap of 
the proposed parental species in bivariate 
space. 



384 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



Table 5. — Factor loadings for the first two principal 
components from an analysis of external measure- 
ments of a pooled sample of males and females in 
definitive alternate plumage of Dendroica striata, D. 
castanea, and their hybrids. 



Principal component axes 



Variable 



II 



Wing chord 

Tail 

Tarsus 

Bill 

Percent variance explained 



0.0076 0.0069 

0.0032 0.0123 

0.0128 0.0025 

0.0130 -0.0095 

40.2% 29.6% 



External Morphology 

As a second step in the diagnosis, hy- 
potheses generated from analyses of plum- 
age characters were tested with analyses of 
morphological size and shape. Size and 
shape characters in birds are assumed to be 
encoded by a multitude of structural and 
regulatory genes, resulting in additive ge- 
netic variation in hybrid morphology. In 
other words, the external dimensions of hy- 
brids should fall within the cumulative 
mensural ranges of characters expressed by 
large samples of the parental species of the 
appropriate age and sex class. 

Univariate comparisons. — External mea- 
surements of the male hybrids fell within, 
or narrowly outside (Texas male) the cu- 
mulative ranges of measurements, for male 
D. striata and D. castanea (Table 1). Mea- 
surements of the wing, tail, and tarsus of 
the hybrids more closely approached those 
of the smaller parental species (D. castanea 
in wing and tarsus, D. striata in tail). In 
fact, the wings and tails of male hybrids 
were more similar in size to females of D. 
striata and D. castanea than to males. 

External measurements of the female hy- 
brid (Table 1) also fell within the range of 
measurements of the hypothesized parental 
species, D. striata and D. castanea. Wing, 
tail, and bill measurements of the hybrid 
were smaller than the character means for 
females of both parental species, while the 
hybrid's tarsi were longer than the parental 
means. 



Multivariate comparisons. — Factor scores 
for male D. striata and D. castanea over- 
lapped extensively along the first two prin- 
cipal components. In bivariate space, the 
male hybrids were closely adjacent to in- 
dividuals of both probable parental species, 
and well inside the 95% confidence ellipse 
(Fig. 10a, Table 5). 

Females of D. striata and D. castanea 
were morphologically less variable than 
males and primarily occupied the lower left 
quadrant of the bivariate plot (Fig. 10b). Fe- 
males of the two species also overlapped 
less extensively in bivariate space than the 
respective males. The female hybrid was 
most similar to D. striata in size and shape. 
Factor scores of the hybrid fell just inside 
the 95% confidence ellipse surrounding the 
centroid of female scores. 

In conclusion, the univariate and multi- 
variate evaluations of external morphology 
and plumage color were consistent with the 
hypotheses generated from the qualitative 
analyses of plumage pattern and color. This 
concordance of results provides strong sup- 
port for the restrictive hypothesis that all 
three specimens represent hybrids of Den- 
droica striata and D. castanea. 

Geographic Range. — As neither Den- 
droica striata nor D. castanea breed in 
southern Michigan, southern Ontario, or 
Texas, the hybrids were presumably mi- 
grating when collected. The bulk of the 
breeding range of D. striata is well north 
of that of D. castanea. However, they over- 
lap extensively in the Northwest Territories, 
British Columbia, Alberta, and Saskatche- 
wan and, to a lesser degree, in the Maritime 
Provinces of Canada (Godfrey 1986, Er- 
skine 1992). The species are essentially par- 
apatric in northern Manitoba (B. Whitney, 
pers. comm.) and in central Ontario (Spier 
1985), approximately due north of the col- 
lection site in Michigan. It is in a region of 
narrow distributional overlap in Ontario and 
Quebec, where population densities of D. 
striata and D. castanea are low (see Spier 
1985), that I expect that hybridization 
would be most likely to occur. These spe- 



VOLUME 109, NUMBER 2 



385 



o 

CL 



0.04 


• 






1 
O ^\ 


1 


- 


0.02 


• 
1 


• 


% 


• 

o 




o 


0,00 






o • 

■ 


o 

• 

o 


• 

o / 


- 


0.02 
nr\A 


\ 

1 




1 


o 

1 


1 





-0.04 



-0.02 



0.00 0.02 

PC 1 



0.04 0.06 



O 

CL 



0.06 



0.04 



0.02 



0.00 



-0.02 - 



-0.04 




-0.04 



-0.02 



0.00 



0.02 



PC I 



0.04 



0.06 



Fig. 10. Bivariate plots of factor scores from a principal components analysis of external morphology (see 
Table 5) for Dendroica striata (empty circles), D. castanea (filled circles), and D. striata X D. castanea hybrids 
(Michigan hybrid = diamond; Texas hybrid = square). Factor scores of males (top) and females (bottom) were 
extracted from the same principal components analysis, but projected on different plots for clarity. 



386 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



cies in first basic plumage are remarkably 
similar and some individuals are difficult to 
distinguish. Juvenile hybrids would be ex- 
traordinarily difficult or impossible to de- 
tect under field conditions. Thus, surpris- 
ingly large numbers of juvenile hybrids 
could migrate unnoticed to their wintering 
grounds each autumn. 

Conclusions 

Hybrid intennediacy in size and shape. — 
Avian offspring from conspecific matings 
usually exhibit high heritability of morpho- 
logical traits (e.g., Boag & Grant 1978, 
Smith & Zach 1979, Smith & Dhondt 1980, 
Dhondt 1982). That is, when the values for 
character traits of the offspring are re- 
gressed on the average of those of the par- 
ents (for monomorphic species), the slope 
of the resulting regression line (= heritabil- 
ity) approaches 1 .0. These data indicate that 
the size and shape of passerine birds are 
largely determined by additive genetic pro- 
cesses, and to a lesser degree, by environ- 
mental factors (James 1983). As such, they 
constitute the empirical foundation upon 
which the morphological diagnosis of hy- 
brid specimens is permitted. 

With a few exceptions (e.g., Maclnnes et 
al. 1989), there have been no comparable 
studies of morphological heritability in avi- 
an hybrids which, strictly speaking, are the 
result of interbreeding of biological species. 
Ornithologists have recognized, however, 
that mensural traits of hybrids were usually 
intermediate between those of their parental 
species long before the field of quantitative 
genetics blossomed (see Falconer 1981). 

But what predictions can be made about 
the morphology of hybrids among distantly 
related species, especially among those 
classified in different genera? If the mixture 
of highly similar genomes results in high 
heritability, then hybridization between spe- 
cies with dissimilar genetics might be pre- 
dicted to result in lower heritability. In oth- 
er words, if heritability, and thus phenotyp- 
ic predictability, is correlated with the de- 



gree of genomic compatibility of the 
parental species, then the phenotypes of 
true intergeneric hybrids are relatively un- 
predictable. Furthermore, the probability 
that hybrids will deviate significantly from 
the biparental midpoint may increase as a 
function of the genetic dissimilarity of the 
parental species (Fig. 11). In operational 
terms, there is no reason to expect a hybrid 
from an intrageneric crossing to occupy a 
position intermediate to the parental species 
in multivariate space. As a corollary, it may 
not be possible to distinguish Fj, F2, or 
back-crosses among genetically divergent 
species, if they do occur, on the basis of 
morphology alone. 

Plumage. — Two assumptions, both 
somewhat vague, are commonly advanced 
in the analysis of hybrid plumage. The first, 
that hybrids express plumage characters of 
both parental species, is bolstered by more 
than a century of careful study of hybrid 
specimens (e.g., Cabot 1854), and finds 
quantitative support in more recent studies 
that indicate that the pattern and color of 
most plumage tracts are polygenically con- 
trolled (see Hutt 1949, Buckley 1982). 

The second assumption is that hybrids do 
not exhibit plumage characters of species 
other than their parents (e.g.. Banks & 
Johnson 1961; Graves 1990, 1992, 1993a, 
1993b; Graves & Zusi 1990). Rohwer's 
(1994) use of "contradictory characters" in 
hybrid evaluation is simply a rephrasing of 
that hypothesis which, in various guises, 
also asserts that luxuriance and atavism do 
not occur in hybrid plumage. As a universal 
statement, the latter is demonstrably incor- 
rect, because "atavistic" characters occur 
with frequency in certain anseriform hy- 
brids (Harrison and Harrison 1963). How- 
ever, atavism has not been adequately doc- 
umented in other avian orders, and plumage 
luxuriance has not been documented in any 
avian hybrid in nature. 

In light of the aforementioned assump- 
tions, perhaps my most noteworthy obser- 
vation on the plumages of the Dendroica 
striata X D. castanea hybrids was the ap- 



VOLUME 109, NUMBER 2 



387 



0) 

o 

0) 
D) 
U 

Q) 4J 
> C! 

■H -H 

73 O 

Pi 

-H "H 

4:1 CO 

M-l CJ 
O 0) 

■P Oi 



-H 



o 



(d M-i 


Oi 



1 , 
1 


Lethal 

- 1 



Genetic difference 
between hybridizing species 



Fig. 11. Hypothetical relationship between the degree of genetic difference between hybridizing avian spe- 
cies and the probability that the morphology of hybrids will significantly diverge from the midpoint of their 
parental species. The dashed line represents the threshold beyond which hybrids die before hatching. 



parent correlation between the degree of 
difference in colorimetric values of homol- 
ogous plumage characters of the parental 
species and the deviation of the hybrid val- 
ues from the averages of the character 
means of the parental species (see Table 2). 
The more discordant the plumage color of 
the parental species, the more likely that 
character in the hybrid closely resembled 
one parental species, rather than a "blend- 
ed" intermediate (e.g., crown color in 
males). This pattern can be partially ex- 
plained by the expansion of possible hybrid 
values when pigmentation of the parental 
species differed markedly. It also suggests 
that a small number of gene complexes con- 
trol pigmentation of individual plumage 
tracts in D. striata and D. castanea. Crown 
pigmentation of D. striata is evidently dom- 



inant (or nearly so) to that of D. castanea 
in hybrids, and as Brodkorb (1934) noted, 
the male hybrid superficially resembles D. 
striata in most respects. Whether this is due 
to genetic dominance or chance recombi- 
nation will only be determined with the dis- 
covery of additional hybrid males. 

Acknowledgments 

I thank Robert Payne and Janet Hinshaw 
(University of Michigan Museum of Zool- 
ogy) for loaning the hybrid specimens, 
Kenneth Parkes (Carnegie Museum of Nat- 
ural History) for forwarding the specimen 
he had borrowed from the San Bernardino 
County Museum, and Carl Hansen for pho- 
tographing specimens. Parkes, Bret Whit- 
ney, and an anonymous reviewer made 
helpful comments on the manuscript. 



388 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



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VOLUME 109, NUMBER 2 



389 



Appendix 1 

Comparative descriptions of the male hybrids, Den- 
droica striata X Dendroica castanea, and their paren- 
tal species in definitive alternate plumage (Figs. 1-4). 
Hybrids are referred to as the "Texas" (SBCM) and 
"Michigan" (UMMZ 53692) specimens, respectively. 

The crown, superciliary, and lores are semi-glossy 
black in striata (Fig. 1). In castanea, the forecrown is 
black becoming into chestnut on the crown above the 
eyes, and extending posteriorly to the rear crown; the 
superciliary and lores are black. The respective parts 
of the Michigan hybrid resemble those of striata; a few 
feathers above the nostrils and on the superciliary have 
faintly buffy-olive tips; several feathers of the hind- 
crown have broader buffy-olive margins. The crown 
of the Texas specimen is more intermediate in appear- 
ance. Feathers of the hindcrown are margined with 
buff-olive, imparting a streaked appearance. Several 
small chestnut spots occur on feathers of the mid- 
crown. 

The parental species and the hybrids have a thin 
nuchal collar: black streaked with white in striata, buf- 
fy-white in castanea, pale buffy-white in the Michigan 
hybrid, and dull olive-buffy in the Texas hybrid. Feath- 
ers on the mantle, scapulars, lower back, rump and 
uppertail coverts of striata have black center stripes 
and broad gray margins. The respective plumage of 
castanea is similarly patterned but the feather margins 
are buffy-olive on the mantle, changing to gray on the 
lower back, rump, and undertail coverts. In the hy- 
brids, dorsal plumage posterior to the hindcrown is 
nearly intermediate in appearance to that of the paren- 
tal species. 

D. striata has a white facial patch that extends from 
the base of the lower mandible, posteriorly to the ven- 
tral eyering and the auriculars. The respective parts of 
castanea are black. In striata, the white facial patch of 
striata is bordered ventrally by a thin black malar stri- 
pe, which originates on the chin and extends to the 
upper breast, and posteriorly by vertical zebra-like 
striping on the nuchal collar. The black "face" of ca5- 
tanea is bordered ventrally by the chestnut throat, and 
posteriorly by a large buff neck patch, a ventral exten- 
sion of the nuchal collar. The facial plumage of the 
Michigan hybrid expresses a complex mosaic of the 
parental characters (Fig. 4). Feathers adjacent to the 
bill are black with pale buffy-white tips; while those 
on the lower eyering and auriculars are grizzled dark 
gray and buffy-white. The auriculars are bordered pos- 
teriorly by a buffy-white neck patch, which is paler 
and smaller than that of castanea. Facial plumage of 
the Texas hybrid is considerably darker than that of 
the Michigan hybrid, and similar in appearance to cas- 
tanea. The posterior auriculars are grizzled (sooty 
black and gray), and bordered posteriorly by a dull, 
silvery-buff, neck patch, which is smaller and less dis- 
tinct than in castanea. 

The remiges and wing coverts of striata and cas- 



tanea are similarly patterned; the greater and middle 
wing coverts are tipped with white in striata but are 
more broadly tipped with pale buffy-white in castanea. 
The hybrids' wings are intermediate in pattern and col- 
or. White spots in the outer rectrices of the hybrids are 
intermediate in size and shape between those of striata 
and castanea. 

The throat, breast, belly, and undertail coverts of 
striata are white bordered laterally by black streaking 
on the sides of the throat, breast, and flanks. The chin 
of castanea is black, bordered posteriorly by chestnut, 
which extends posteriorly to the upper breast along the 
midline and laterally to the flanks; the center of the 
breast, lower belly, and undertail coverts are buffy- 
white. The feathers of the Michigan hybrid's throat are 
grizzled with black and chestnut, many faintly tipped 
with white, especially on the chin and along the mid- 
line, imparting the appearance of wide malar stripes. 
The breast, belly, and undertail coverts of the Michi- 
gan hybrid are white, faintly tinted with buff; the 
flanks are marked with muted brownish-black spots 
and streaks and a strong chestnut wash. The chin, 
throat, upper breast, and sides of the Texas hybrid are 
"roan" (sooty black, subtly grizzled with chestnut and 
pale gray) in a pattern mirroring the distribution of 
chestnut in castanea. Center of the breast, belly and 
undertail coverts of the Texas specimen are pale buffy 
white. A few indistinct pale chestnut and sooty spots 
are found on the flanks. 

Appendix 2 

Comparative descriptions of the female hybrid 
(UMMZ 216628), Dendroica striata X Dendroica cas- 
tanea, and its parental species in definitive plumage 
(Figs. 5-7). 

Feathers of the crown, scapulars, and mantle of 
striata are olive to grayish-olive with pronounced 
blackish-brown shaft streaks. The respective plumage 
of castanea is olive-gray to gray but more heavily 
streaked with black; feathers of the crown, from the 
eyes to the nape, are chestnut distally with grayish- 
olive margins. The crown of the hybrid is more heavily 
streaked than in striata, but less so than in castanea; 
feather margins are olive (a few grayish-olive) and ex- 
hibit no trace of chestnut pigmentation on the basal or 
distal barbs under magnification (7X). The mantle of 
the hybrid, which contrasts with its olive crown, is 
intermediate in appearance between those of the pa- 
rental species. The rump and uppertail coverts are ol- 
ive-gray in striata and gray in castanea. The hybrid 
has an olive-gray rump and gray uppertail coverts. 

The short superciliary of striata is poorly defined by 
pale olive to pale yellow feather tips. In castanea the 
superciliary is even more poorly defined and streaked 
with dark gray and black. The lores and auriculars of 
striata are mottled olive-gray, bordered posteriorly by 
a faintly perceptible olive-gray patch on the side of the 
neck. The lores and auriculars of castanea are more 
heavily pigmented and contrast with the throat and a 



390 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 



well-defined buff neck patch. Pigmentation of the su- 
perciliary, lores, and auriculars of the hybrid falls with- 
in the range of variation found in striata, but the au- 
riculars are bordered posteriorly by well-defined buffy 
neck patch as in castanea. 

White tips of the middle and greater wing coverts 
of the hybrid are intermediate in width between those 
of striata and castanea. Outer margins of the rectrices 
are olive-gray in striata and the hybrid and gray in 
castanea. 

The venter of striata is white tinted with pale yellow 



or olive on the throat and breast; narrow black streak- 
ing occurs on the side of the throat, breast, and flanks. 
The venter of castanea is buffy-white to buff, exten- 
sively suffused with chestnut on the throat, side of the 
breast, and flanks. The throat, upper breast, and flanks 
of the hybrid are buff, fading to pale buffy-white on 
the belly and undertail coverts. Several flank feathers 
are marked with pale chestnut and faint muted streaks 
appear at the side of the breast. The dried tarsi of the 
hybrid are intermediate in color between those of stria- 
ta (light brown) and castanea (dark brown). 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

109(2):391-396. 1996. 

Early record of indricothere (Mammalia: Perissodactyla: 
Hyracodontidae) from the Aral Sea region of western Kazakhstan 

Spencer G. Lucas and Robert J. Emry 

(SGL) New Mexico Museum of Natural History and Science, 1801 Mountain Road N.W., 

Albuquerque, New Mexico 87104, U.S.A.; 

(RJE) Department of Paleobiology, National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 20560, U.S.A. 

Abstract. — Upper molar fragments of the indricothere rhinoceros Paracer- 
atherium sp. are described from the Chilikta Formation at Altyn Chokysu, north 
of the Aral Sea in western Kazakhstan. Marine bivalves indicate the Chilikta 
Formation is of early Oligocene (Late Rupelian) age, and thus provide the first 
direct cross-correlation of an occurrence of Paraceratherium with the marine 
timescale. This find extends the temporal range of Paraceratherium in the Aral 
Sea region back from the late Oligocene to the early Oligocene, making it 
consistent with the temporal range noted for the genus in China. Paracera- 
therium thus had a geologically synchronous first appearance across Eurasia 
during the late early Oligocene. 



The largest land mammals of all time 
were the indricotheres, giant rhinoceroses 
that lived during the early to middle Ceno- 
zoic in Eurasia. Indricothere evolution began 
during the middle Eocene with the pony- 
sized genus Forstercooperia and culminated 
during the Oligocene-early Miocene with 
Paraceratherium (=Baluchitherium, =In- 
dricotherium), a rhinoceros that stood more 
than five meters tall at the shoulder, the larg- 
est land mammal of all time (Granger & 
Gregory 1936, Lucas & Sobus 1989). 

One of the most important collecting ar- 
eas for fossils of Paraceratherium is north 
of the Aral Sea in western Kazakhstan (Fig. 
1). Here, the most nearly complete skeleton 
known of the genus was collected (Orlov 
1939, Gromova 1959) from the Kumbulak 
Cliffs east of the town of Agyspe (Akespe) 
along Perovsky Bay on the northern shore 
of the Aral Sea. This and other specimens 
of Paraceratherium from the region north of 
the Aral Sea are restricted to the Aral For- 
mation, strata of late Oligocene (Russell & 
Zhai 1987) or early Miocene (Akhmetyev & 
Sychevskaya 1994) age. Here we document 



a much older occurrence of Paracerather- 
ium in this region that clarifies the temporal 
distribution of the genus. In this article, 
AMNH refers to the Department of Verte- 
brate Paleontology, American Museum of 
Natural History, New York; and USNM re- 
fers to the Department of Paleobiology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 

Stratigraphy and Provenance 

North of the Aral Sea, four rock-strati- 
graphic units of Eocene-Oligocene are ex- 
posed (Akhmetyev & Sychevskaya 1994) 
(Fig. 2). The oldest, the Chegan Formation, 
is yellowish green, bentonitic marine shale 
with dinocysts and molluscs of late Eocene 
(Priabonian) age. The Kutanbulak Forma- 
tion disconformably overlies the Chegan, is 
as much as 28 m thick and is composed of 
mostly yellow, orange and brown, fine- 
grained quartzarenite. It is unconformably 
overlain by the Chilikta Formation, as much 
as 23 m of shale and thinly interbedded 
shale-sandstone. The Chilikta Formation 



392 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASfflNGTON 



Miles 


100 


200 




300 


Kilomatefs 


100 200 


300 


400 


500 



Russia 




r'' 



Novosibirsk 




■ -/T'etropavlovsk'. . 



yl 



Aktyubinsk 



Kazakhstan 

Aral -^^Altyn-Chokysu 

Sea 




\ 

Semipaiatinsk 

Lake ^S^ 
Zaysan 

Ayaguz p 

mj 

Lake ■ 

Balkhash ^^' 



H 



^ 




Kzyl-Orda 



. / ■ /C" * ->^ Uzbekistan ^ 



•^ 



-^ 



Turkmenistan 



\ 



\ 



\ 



Alma-Ata • < 

'. Tashkenty.'^ Kyrgyzstan ^ 

.-Tc'-XC^-^ China 

Tajikistan ^. 



/,. 



^ 



Iran 



C--^ ixj^;^- 



I Afganistan ^' Pakistan 



Fig. L Map of Kazakhstan showing location of Altyn Chokysu north of the Aral Sea. 



produces marine bivalves (especially Er- 
genica cymlanica) that correlate it to the 
Solenovo horizon of the Crimea-Caucasus 
of late Early Oligocene (late Rupelian) age 
(Akhmetyev & Sychevskaya 1994). 

The Chagray Formation disconformably 
overlies the Chilikta Formation and is as 
much as 33 m thick and mostly yellowish 
gray and brown, micaceous sandstone. The 
Aral Formation conformably overlies the 
Chagray. It is at least 24 m thick north of 
the Aral Sea (its top is everywhere eroded) 
and consists mostly of grayish yellow 



green, calcareous shale and claystone char- 
acterized in its lower part by numerous 
lenses of the euryhaline bivalve Corbula. 
All fossil mammal localities reported by 
previous workers (Russell & Zhai 1987, 
Bendukidze 1993, Akhmetyev & Sychev- 
skaya 1994) from north of the Aral Sea are 
in the Aral Formation. We follow Russell 
& Zhai (1987) in regarding these mammals 
as late Oligocene in age, but Soviet scien- 
tists regard them as early Miocene in age 
(Akhmetyev & Sychevskaya 1994). 

The new locality at which we collected 



VOLUME 109, NUMBER 2 



393 



LU 

Z 
LU 

o 
o 

_J 

o 



LU 

I- 
< 



> 

_J 

DC 
< 
LU 



LATE 
EOCENE 



ARAL FORMATION 



previous records of Paraceratherium 

CHAGRAY FORMATION 

— — — disconformity— — — — 



Paraceratherium reported here 

CHILIKTA FORMATION 

KUTANBULAK FORMATION 

— — -- disconformity— — — — — 



CHEGAN FORMATION 



Fig. 2. Summary of Eocene-Oligocene stratigraphic units in Altyn Chokysu area showing stratigraphic levels 
of Paraceratherium occurrences (lithology schematic). 



Paraceratherium tooth fragments is in the 
Ckilikta Formation on the escarpment of 
Altyn Chokysu, a plateau about 20 km 
northwest of the town of Saksaulskaya and 
about 70 km northeast of the Kumbulak 
Cliffs (Figs. 1-2). This locality is at UTM 
357737 IE, 5238795N, zone 41, stratigraph- 
ically well below Bendukidze's (1993) 
"Shokysu" mammal locality in the Aral 
Formation. The fossiliferous horizon is a 
1.2-m-thick, white, fine-grained sandstone, 
10.8 m above the base of the Chilikta For- 
mation, which is 14.4 m thick at this loca- 
tion. The Paraceratherium tooth fragments 
were associated with a sirenian rib, fish 
bones and teeth of the sand tiger shark Car- 
char ias sp. 

Systematic Paleontology 

Order Perissodactyla Owen 1848 

Family Hyracodontidae Cope 1879 

Subfamily Indricotheriinae Borisyak 1923 

Genus Paraceratherium Forster Copper 

1911 



Paraceratherium sp. 
Fig. 3 

Referred specimen. — USNM 482243, ap- 
proximately 50 fragments of cheek teeth, 
three of which, illustrated here, are parts of 
ectolophs of right M', M^, and M^. 

Description. — The three largest tooth 
fragments, here referred to as 1, 2, and 3, 
are portions of upper molar ectolophs. Frag- 
ment 1 is right M^ ectoloph from paracone 
through parastyle, and part of protoloph. 
Paracone forming thick rib on labial face of 
ectoloph and separated from parastyle by 
distinct cleft. Parastyle a more prominent 
and anteriorly-projecting rib. Ectoloph 
making sharp, nearly right-angled fold and 
becoming confluent with protoloph. Mini- 
mum crown height at paracone 59 mm. 

Fragment 2 from anterior part of ecto- 
loph of right M-. Preserves paracone as a 
less prominent rib than on M^ ectoloph. 
Cleft between paracone and parastyle less 
pronounced than on M^. Paracone crown 
height about 66 mm. 

Fragment 3 part of posterior portion of 



394 



PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASmNGTON 




Fig. 3. Tooth fragments of Paraceratherium sp. from Altyn Choky su compared to cast of holotype of 
Paraceratherium asiaticwn. A-C, USNM 482243, Paraceratherium sp. from Altyn Chokysu: A, occlusal view 
of fragment of right M^ ectoloph (fragment 1 in text); B-C, occlusal view of right M^ ectoloph (fragments 3 
and 2, respectively, in text). D-E, AMNH 26972, cast of holotype of Paraceratherium asiaticum, left M'-^ 
(photographs reversed): D, occlusal; E, labial views. F-H, USNM 482243: F, anterior view of fragment of right 
M- ectoloph (fragment 1 in text); G-H, labial views of fragments of right M^ ectoloph (fragments 3 and 2, 
respectively, in text). 



right M- ectoloph, probably from same 
tooth as fragment 2, but two pieces do not 
fit together. Blade-like metacone projecting 
from rest of ectoloph occlusal edge. Prom- 



inent basal cingulum on posterior edge of 
crown. Fragment 2 and 3 suggest total ec- 
toloph length of more than 90 mm. 

Identification. — We compared these 



VOLUME 109, NUMBER 2 



395 



tooth fragments to casts of upper dentitions 
of Paraceratherium asiaticum from the 
Turgay region of central Kazakhstan (Fig. 
3: Granger & Gregory 1936, figs. 2C-D). 
These casts are AMNH 26971, left P^-M^, 
and AMNH 26972, right P^-M^ (cast of ho- 
lotype of P. asiaticum). They are very sim- 
ilar in size and morphology to the frag- 
ments we collected at Altyn Choky su. Frag- 
ment 1 has a minimum crown height of 59 
mm